June 1999 North American Native Orchid Journal

NORTH AMERICAN
NATIVE ORCHID JOURNAL

______________________________________
Volume 5 June
Number 2 1999
a quarterly devoted to the orchids of North America
published by the
NORTH AMERICAN
NATIVE ORCHID ALLIANCE
* * * * * * *
* * * * * * *
IN THIS ISSUE:
PROCEEDINGS OF THE 4th ANNUAL NORTH
AMERICAN NATIVE ORCHID CONFERENCE
THE UNPREDICTABLE WEATHER
POLLINATOR OF CLUSTERED LADYS-SLIPPER
ORCHID IN OREGON
A STRATEGY FOR COPING WITH RARITY IN THE
ORCHID CYPRIPEDIUM FASCICULATUM….and more
NORTH AMERICAN NATIVE
ORCHID JOURNAL
(ISSN 1084-7332)
published quarterly in
March June September December
by the
NORTH AMERICAN NATIVE ORCHID ALLIANCE
a group dedicated to the conservation and promotion of our
native orchids
Editor:
Paul Martin Brown
Assistant Editor: Nathaniel E. Conard
Editorial & Production Assistants:
Philip E. Keenan
Stan Folsom
Nancy Webb
The Journal welcomes articles, of any length, of both a scientific

and general interest nature relating to the orchids of North
America. Scientific articles should conform to guidelines such as
those in Lindleyana or Rhodora. General interest articles and notes
may be more informal. Authors may include line drawings
and/or black and white photographs. Color inserts may be
arranged. Please send all inquiries or material for publication to
the Editor at PO Box 772121, Ocala, FL 34477-2121 (late May -
early Oct. Box 759, Acton, ME 04001-0759).
2000 Membership in the North American Native Orchid Alliance,

which includes a subscription to the Journal, is $26 per year in the
United States, $29US in Canada and $32US other foreign
countries. Payment should be sent to Nancy A. Webb, 84 Etna
St., Brighton, MA 02135-2830. Claims for lost issues or canceled
memberships should be made to the editorial office within 30
days.
NORTH AMERICAN NATIVE
ORCHID JOURNAL
Volume 5 June
Number 2 1999
CONTENTS
NOTES FROM THE EDITOR
75
The Slow Empiricist
77
4TH ANNUAL NORTH AMERICAN NATIVE ORCHID
CONFERENCE
Philip E. Keenan
84
ORCHIDS AT RABENAU CAMP
Chuck McCartney
90
PLATANTHERA HYPERBOREA AND A
REAPPRAISAL OF SOME BASIC GREEN
PLATANTHERAS
Charles J. Sheviak
115
ORCHIDS OF MEXICO
Eric A. Christenson
142
PHYLOGENY AND BIOGEOGRAPHY OF
POGONIINAE (VANILLOIDEAE: ORCHIDACEAE)
Kenneth M. Cameron
151
KEYING OUT A RED MITSUBISHI
or the Trials, Travails, and Tears of an Orchid Hunter
Tom Sampliner
164
RECENT TAXONOMIC AND DISTRIBUTIONAL
NOTES FROM FLORIDA 2.
A Striking New Color Form of Sacoila lanceolata
Paul Martin Brown
169
HOW IT ALL CAME ABOUT
Stanley N. Folsom
174
POLLINATOR OF CLUSTERED LADYS-SLIPPER
ORCHID, CYPRIPEDIUM
FASCICULATUM, (ORCHIDACEAE) IN OREGON
C. S. Ferguson & K. Donham
180
A STRATEGY FOR COPING WITH RARITY IN THE
ORCHID CYPRIPEDIUM FASCICULATUM
Penelope A. Latham
185
LOOKING FORWARD:
September 1999
inside back cover
Color Plates:
1. p. 197 Habenaria quinqueseta
2 p. 198 Basic Green Platantheras
3. p. 199 Sacoila lanceoalta var. lanceolata forma folsomii
4. p. 200 Cypripedium. fasciculatum
Unless otherwise credited, all drawings in this issue are by Stan Folsom
Unless otherwise credited graphics (charts, maps etc. ) are created by the
individual authors.
The opinions expressed in the Journal are those of the authors. Scientific
articles may be subject to peer review and popular articles will be examined for
both accuracy and scientific content.
Volume 5, number 2, pages 75-200; issued June 24, 1999.
Copyright 1999 by the North American Native Orchid Alliance, Inc.
Cover: Cleistes bifaria by Stan Folsom
NOTES FROM THE EDITOR
With this the June issue for 1999 I trust the

Journal is now back on track. If you are still missing
any issues, please let me know so they can be replaced.
Several changes are being made in the Journal - all of
which I truly hope will improve both the quality and
service.
Because of the labor involve, and the lack of any
help assembling the Journal, all of the color will
appear as the last set of pages in each issue.
Starting with this issue all mailings will be by Book
Rate rather than by Bulk Mail. We are losing too
many issues with the Bulk Mail and the service has
been unreliable.
The subscription rate for the Journal will not change,
although the cost of postage will increase
considerably. I will make every effort not to
decrease the annual size of the Journal. Several
members have already offered to include extra with
the renewals to help cover the additional postage -
this is most appreciated.
Foreign mailing to Canada and Europe will be
greatly improved by sending all of the Journals to
one address and then they will be mailed out with
that area from there. We have a member in Canada
and one in England who have offered to assist with
this.
75
There are many interesting and exciting articles in this
issue, not all of which are from the 1999 conference. I
am especially pleased with the pair of article on
pollinators and rarity of Cypripedium fasciculatum from
Oregon. Although they are brief, please take time to
read them carefully.
Plans are well underway for the 5th Annual North
American Native Conference to be held in the majestic
Olympic Mountains and Cascade Range of Washington
in July of 2000. Registration information is within this
issue.
The September issue will be printed and assembled in

late September when we return to Florida
Paul Martin Brown

Editor
PO Box 759,
Acton Maine 04001
207/636-3719
naorchid@aol.com
after September 15:

PO Box 772121
Ocala, Florida 34477
352/861-2565
76
Empiricist: THE UNPREDICTABLE WEATHER
The Slow Empiricist
It seems that the weather patterns change almost

as often as years go by. This variability then affects the
growing and blooming times of all natural floras in a
given area. Controlled conditions in the home or
greenhouse can overcome much of the vagaries that an
ordinary year or two of weather can bring to the
outdoors. In a controlled situation, you can compensate
for long periods of overcast skies with lighting
conditions. Periods of drought or prolonged wet spells
won't affect indoor specimens that have carefully
regulated watering times and amounts. This is not so in
nature, however, and it can produce a variety of results.
My first year in retirement in Florida in 1997 was

marked by periods of heavy and frequent rain most of
the winter season. I was beginning to wonder if Florida
was aptly nicknamed, The Sunshine State, we had so much
inclement weather. Many areas were threatened with
flooding. The weather people blamed the conditions on
El Nino, and excessive warming of part of the Pacific
waters that seemed to alter the normal rhythms of the
jet stream. That was followed by a hot and very dry
77
summer, which produced many wild fires that burned

over 700,000 acres of land in the state. The fall season
continued the drought and it has persisted into the
spring of 1999. The current weather has been hotter
than normal as well. According to the weather experts
La Nina has replaced El Nino and we can expect more
drought conditions until it abates.
How has this impacted the world of native

orchids? It has been a curious mix of events. The North
American Native Orchid Alliance's annual conference
was held in Florida in the month of April. Specifically, it
was held April 8, 1999 to April 13, 1999. There were
field trips set up to catch as many species in bloom as
possible. The weather was not cooperating. Florida
ladies'-tresses, Spiranthes floridana was blooming early
and going to fruit. In 1998, it was holding bloom well
into the middle of April. Eaton's'-tresses, Spiranthes
eatonii, a newly described species1 was not even showing
itself in the Jennings Forest State Forest were there
were several dozen plants last year. The short-lipped
ladies'-tresses, S. brevilabris was also blooming earlier
but it at least had several plants that would hold for the
group that hoped to take photographs. To compound
the problems, many of the blooming plants of these
Spiranthes were smaller than usual.
The prolonged dry spell and the warmer than

normal conditions had pushed the plants into a seeming
rush-mode or a non-mode of blooming. Green
1
See article by P.M. Brown in March 1999 issue of NANOJ p. 3-15.
78
adder's-mouth, Malaxis unifolia, which was abundant in

1998, had not even shown itself by April 8, 1999,
joining those orchids that chose the non-mode style of
bloom. The search for it at the SR 200 site in Ocala
turned up a new species for that particular area, though.
English friends, Mike and Carol Parsons, while
exploring the area found the speckled ladies'-tresses,
Cyclopogon cranichoides, blooming in late March. They
were smaller than the ones at Alexander Springs
Recreation Area on the other side of Ocala but they
were there and gave the state a fourth extant for the
species.
Paul Martin Brown, who organized the

conference, spent many hours scouting the projected
field trip sites and scoured the territory to find viable
plants for the group. Contingency plans changed as the
plants followed their own paths for bloom. The group
ended up going into southern Georgia to observe
Spiranthes eatonii. It was in spike there and just coming
into flower. They were also able to see bearded grass-
pink, Calopogon barbatus and pale grass-pink, pale grass-
pink, C. pallidus, along the roadsides on the way to the
S. eatonii site.
Just the week before, in preparation for the

conference field trip there, Paul and I were in the
Jennings State Forest scouting for small spreading
pogonia, Cleistes bifaria, and Spiranthes eatonii, when I
found several plants of the many-flowered grass-pink,
C. multiflorus. Two were only in bud. When Paul went
back the following Tuesday, two days before the
79
scheduled field trip as a last minute check, they had all

gone to fruit.
Weather has certainly played a major factor in

man's plans for viewing orchids or for that matter, any
plant. Being flexible helps a person weather the vagaries
that effect flowering time. Unfortunately in this day and
age man can do very little to alter weather conditions. It
is probably a bad idea to even venture to try, as nature
seems to know what its doing even if we don't
particularly care for the current status of things. The
wild fires of last year are a good example of how good
can come from a bad situation. This is not to trivialize
the loss of property and income that these fires created
but many Florida orchids are respondant to fire as a
means of flowering. Calopogon multiflorus, Cleistes bifaria
and the large spreading pogonia, C. divaricata all
appear in greater numbers in areas that have been
burned over in the year previous. In May we found new
sites for both species of Cleistes, one is in Cary State
Forest and the other in the Raiford Wildlife
Management area, where we had only found C.
divaricata last year. Fire, if it is not all consuming, seems
to cut back the competition, open up the forest floor
and allow these plants to flourish.
My best advice is to persist in your quests and

adjust to peculiarities of nature that you find yourself in,
in any given year. Fretting over a current condition that
is frustrating can be alleviated if you consider that this
gives you an opportunity to revisit this site on a more
favorable year and season. The Slow Empiricist
80
many-flowered grass-pink
Calopogon multiflorus
81
pale grass-pink
Calopogon pallidus
82
small spreading pogonia

Cleistes bifaria
83
Keenan: 4th ANNUAL NORTH AMERICAN NATIVE ORCHID
CONFERENCE
4TH ANNUAL NORTH AMERICAN

NATIVE ORCHID CONFERENCE
Philip E. Keenan
Although the University of South Florida in Tampa,

Florida hosted the weekend of meetings during the 4th
Annual North American Native Orchid Conference, it would
be fair to say that the four days of field trips ranged all
the way from the Georgia border to the Big Cypress
region of extreme south Florida. As usual at these
annual meetings, Tampa featured some of the biggest
names in the orchid world.
John Beckner of the Orchid Identification

Center, Marie Selby Botanical Gardens, led things off
with a provocative talk entitled Florida - the Rules are
Different Here!, emphasizing some of the extremes of the
state: hurricanes, droughts, fires, "terminal" water
supply situation, over-development, over-population
(now the fourth largest state in population!, behind only
New York, Texas and California), illegal trade in drugs,
plants and people, and the dangers of attempting the
Fakahatchee Swamp alone. If all this wasn't bad enough,
and if the pollinators go!
84
CONFERENCE
Roger Hammer of the Dade County Park Service

followed with a slide presentation on the Orchids of South
Florida, especially in the same Fakahatchee Strand,
featuring a marvelous slide of the incredible night-
fragrant epidendrum, Epidendrum nocturnum, and his
favorite of all, the ghost orchid, Polyradicion lindenii.
Charles J. Sheviak of the Biological Survey, New York
State Museum, attempted to ascertain, explain, and
rationalize some of the uncertainties that plague
Platanthera hyperborea and other green platantheras that
most of us prefer to just ignore as a "hybrids of what"
problem as he so aptly put it.
Larry Zettler, a biologist at Illinois College,

presented an update on the symbiotic seed germination
of eastern prairie fringed orchis, Platanthera leucophaea,
yellow fringeless orchis, P. integra, and northern oval
ladies'-tresses, Spiranthes ovalis var. erostellata with his
usual precise graphics and impeccable speech. Paul
Martin Brown filled us in on "a new name for an old
discovery" a reference to Eaton's ladies'-tresses,
Spiranthes eatonii, in his usual professional and forthright
manner.
On Sunday, Eric Christenson, a Ph.D. research

scientist in Florida indulged in a change of pace with an
erudite review of the Orchids of Mexico. Chuck
McCartney, former editor of the AOS Bulletin, followed
with an excellent slide show on the Orchids of Rabenau
Camp featuring everyone's favorite Polyradicion lindenii,
the sensational ghostly flying frog of folklore. His three
slides of this remarkable orchid were easily "the best in
85
CONFERENCE
show." Wendell Vaught and Mike Owen, both of the

Florida Park Service, teamed up with a history of the
Fakahatchee Strand.
Two members presented some of there favorites

with Dennis Horn showed his slides of a trip with Will
and Hazel Caster to Götland, Sweden and the
abundance of orchids to be found there while Dennis
Maleug shared some of his striking photos of the
various fringed orchids.
Ken Cameron of the New York Botanical

Garden, spoke on his specialty, the Phylogeny and
Biogeography of Pogoniinae, the bottomline for which is still
based on educated guesses, in terms of "how did they
get where they are now," for one example, but we are
still trying to put the puzzle together.
The concluding speaker was Robert Dressler who

"regaled" us with The Theory and Practice of Classification,
relating how several individual plant characters have
been termed the "best" by many different taxonomists
over the centuries, in determing species, but in his
opinion, a variety of characters are the key in the final
analysis. Incidentally, Dr. Dressler had the best joke of
all the speakers; one scientist to the other: "what do you
think of God?" "Well, he must have had a great
fondness for beetles!" (largest number of species of any
plant or animal --- as we all knew, right?)
Mike Owen, biologist for the Fakahatchee Strand

State Preserve deservedly received the 1999 Alliance
86
CONFERENCE
Conservation & Education Award for his meticulous

record keeping and generous sharing of his knowledge
of the native orchid world with his devoted work at the
Fakahatchee Strand.
Two days of field trips before the two-day

weekend meetings were followed by two more days on
Monday and Tuesday. Because of severe drought
conditions in all of Florida for several weeks prior to the
second week of April conference dates, the scorecard of
orchids in bloom fell short of the norm. Nevertheless,
there were several excellent orchids in prime condition
to satisfy most of us, even though it required traveling
the first day up to St.George in Georgia (where the state
line does a little belly flop into Florida just west of
Jacksonville). Here we picked up three grass-pinks -
bearded grass-pink, Calopogon barbatus, common
grass-pink, C. tuberosus, and pale grass-pink, C. pallidus
in the same roadside "bog" which was perfectly dry on
this day. Lesser pitcher plant, Sarracenia minor, the most
common pitcher-plant in Florida, and parrot pitcher-
plant, S. psittacina, whose leaves lie nearly flat on the
ground, were bonus plants.
Back in Florida, near Middleburg and Starke we

added the very rare and tiny Florida ladies'-tresses,
Spiranthes floridana and Eaton's ladies'-tresses, S.
eatonii, the latter reminding of an even smaller southern
slender ladies'-tresses Spiranthes gracilis. The premier
orchid for me turned out to be the diminutive speckled
ladies'-tresses, Cyclopogon cranichoides, in peak bloom at
the Alexander Springs Recreation Area in the Ocala
87
CONFERENCE
National Forest. A hand lens made the difference on

this one as it does with so many of our native orchids.
West of Ocala, the short-lipped ladies'-tresses,
Spiranthes brevilabris was in good bloom, apparently on
only one road shoulder.
The excessive heat of an early Spring knocked

out a remote cemetery site for the bronze green-fly
orchid, Epidendrum conopseum var. mexicanum. But further
south near Avon Park, we hit the jackpot with hundreds
of prime water spider orchids, Habenaria repens, prolific
along the water-less ditches. In one of the dry ditches
several plants of the white-lipped form of early ladies'-
tresses, S. praecox forma albolabia, were flowering. On
the dry road shoulders above these same ditches were
hundreds of grass-leaved ladies'-tresses, Spiranthes
vernalis, also in prime condition.
In the Disney Preserve near Kissimmee, we

caught the last of the many-flowered grass-pink,
Calopogon mutiflorus, in bloom along with a couple of
rattlesnakes for good measure, putting to good use the
walking sticks we carried on the trip, much to the
puzzlement of the non-believers who passed us by.
Spiranthes praecox - without the green stippling - (forma
albolabia) waved in the breezes along equally dried up
roadsides not too far out of the Disney Preserve.
At Highlands Hammock State Park, we saw the

same colony of jingle bell orchid, Harrisella porrecta,
which Luer used in his classic volume of the nineteen
seventies, in the expected fruiting stage, supposedly
88
CONFERENCE
superior to the flowers. A big disappointment for me

was the absence of any Sacoila lanceolata, the red ladies'-
tresses, which I had remembered on my only other trip
to Florida more than forty years before!
Our last day, after a brief stop at the airport by

the visitors center for the Big Cypress Preserve, where
we saw Simpson's grass-pink, Calopogon tuberosus var.
simpsonii, featured the long anticipated Fakahatchee
Swamp and everything the folklorist had conjured up in
our minds over these many years of anticipation. But
alas, the drought had pretty much dried up even this
bastion of the subaqueous. We did see one dingy star
orchid, Epidendrum amphistomum, in full flower with its
dingy coppery flowers. However, rather than harbor
feelings of defeat and dismay, we went away, knowing
that the future would surely present an opportunity to
experience this wilderness kingdom in all its glory.
Philip E. Keenan, 31 Hillcrest Dr., Dover, NH. 03820

sk2325@server1.rscs.net
Phil has written frequently for the journal and annually recaps the
conferences for us. He is the author of the recently published
Wild Orchids Across North America published by Timber Press.
89
McCartney: ORCHIDS AT RABENAU CAMP
Chuck McCartney
In March of 1979, while on a botanical tour in

Colombia, I met a man named Bill Rabenau and his 16-
year-old son, William. They lived in Davie, Florida, a
western suburb of Fort Lauderdale in Broward County,
the same county where I lived and worked.
We got to be friends, and in September of that

year, Bill invited me to spend the weekend at his
hunting camp in the Big Cypress Swamp. Although I
am a third-generation South Floridian, I had spent
relatively little time in the Big Cypress when Bill
extended his invitation, so I wasn't quite sure what to
expect. As it turned out, I found something rather close
to paradise for an amateur botanist/naturalist.
The word "camp" made me envision rather crude

living conditions. But Bill's place had most of the
comforts of home -- minus the annoyance of a
constantly ringing telephone.
But even better than the accommodations was the

setting. Bill's main camp occupied a small oak-shaded
tree island surrounded by a rich mosaic of plant
90
communities that included pinelands, open glades,

cypress forests, and, best of all to my orchid-loving soul,
deep, mysterious swamps of pond apple, Annona glabra
and pop-ash, Fraxinus caroliniana.
Although this central camp encompassed only 90

acres, Bill also owned about 20 acres of mostly pineland
converted to pasture about a half-mile to the east and
50 acres of mixed hammock, pineland and swamp a
mile to the west of his main property. In my mind, this
whole area came to be known as Rabenau Camp, and I
still use that designation for the region.
Rabenau Camp is in the northeastern Big Cypress

Swamp in Collier County, Florida, near where it comes
together with Broward and Hendry counties. Being at
the edge of the Big Cypress, the area seems to have
more varied plant communities than are encountered
deeper into this vast tract of wilderness making up
much of the southwestern tip of the Florida Peninsula.
The diversity of habitats makes for a rich and varied
flora, and that diversity extends to the orchids of the
area.
In the company of Bill's son, William (who was,

like this father, an excellent woodsman), I started to
explore the fascinating area around Rabenau Camp.
Later, when William went away to college, I continued
these explorations on my own.
In the nearly 20 years that I have been botanizing

in the area of Rabenau Camp, William and I have
91
encountered 29 species of orchids. All of these species

occurred in an area of only about six square miles. This
is more than one-fourth of the orchids reported for
Florida and it represents one of the highest
concentrations of orchid species to be found anyplace
in North American north of Mexico. The only possible
contender for this distinction might be the fabled
Fakahatchee Strand, which lies along the opposite
western border of the Big Cypress from Rabenau Camp.
The orchid list for the Fakahatchee (Austin, undated)
numbers 45 taxa (some questionable). But the species
there are spread out over a geographic area of more
than 126 square miles. Still, there may be some six-
square-mile area within the Fakahatchee that could
harbor as many species as are found in the region of
Rabenau Camp.
Of the 29 species of orchids discovered in the

vicinity of Rabenau Camp, 11 are epiphytes. This
number has remained constant since our first years of
exploration in the area. However, the number of
terrestrial species discovered keeps growing, with the
latest two species recorded as recently as June 1996 and
June 1997. To date, the number of terrestrial orchid
species observed in the Rabenau Camp area stands at
18.
The following is a quick look at the orchids

discovered in this fascinating area of southern Florida.
The nomenclature is conservative, with proposed name
changes discussed where necessary:
92
THE EPIPHYTES
Epidendrum anceps: A fairly common species of

the pond apple swamps, where some plants grow to be
quite large. A few plants produce reddish-purple rather
than green leaves. A winter/spring bloomer with
hemispherical clusters of small flowers, ranging in color
from bronzy brown to green. Reblooms from the same
inflorescence. Epidendrum monographer Eric Hagsater
of Mexico has proposed the name Epidendrum
amphistomum for these Florida plants (personal
communication), but I can't see how he distinguishes
our plants from Epidendrum anceps.
Epidendrum difforme: Rare in this section

of the Big Cypress. Found in cypress swamps and
adjacent pond apple/pop-ash sloughs. Blooms in
August and September with umbellate clusters of
translucent concolor chartreuse flowers. The taxonomy
at present is very confused for these plants. The
segregation into the genus Neolehmannia has pretty
generally been rejected. But Hagsater (1993) says the
Florida plants are different from the type form of
Epidendrum difforme, so he has renamed our plants
Epidendrum floridense.
Epidendrum nocturnum: Relatively common in

the pond apple swamps of the region, with some plants
growing to nearly three feet tall. Flowers in fall/winter,
producing surprisingly large, although spidery, flowers
with greenish-brown sepals and petals and a glistening
white, three-lobed lip with two bright yellow calluses at
93
the entrance to the tube formed by the adnation of the

lip and column. Typical of many Florida plants of this
species, plants at Rabenau Camp most often produce
cleistogamous flowers, meaning they fertilize themselves
within the maturing bud and go directly to seed without
ever opening. This doesn't seem to be entirely genetic,
though, because plants with cleistogamous flowers
occasionally will produce a flower that opens normally.
Epidendrum rigidum: A sprawling rhizomatous

miniature species, which sometimes forms large
colonies. Fairly common at Rabenau Camp. It is, in
truth, the kind of orchid that only an orchid lover could
love. The tiny green flowers appear most commonly in
the fall and winter months, although they may be found
sporadically at other times.
Encyclia tampensis: The only Encyclia in the
strictest sense verifiably reported for Florida2. Once
common in the Rabenau Camp area, where it grew in
habitats ranging from sunny, open forests of pond
cypress, Taxodium ascendens, to tree island hammocks and
pond apple swamps. Seems less frequent in the area
now, with many dead plants found attached to the trees.
Flowers primarily from late spring through summer,
producing flowers with a honey-sweet fragrance, mainly
in the afternoon. Flower color is variable, but mostly
with brownish-yellow sepals and petals and a white lip
2A specimen of Encyclia rufa collected by Small from 'near Eau Gallie' Brevard
County has been found at the New York Botanical Garden. No other record
or report of this species has ever been found for Florida. It must remain a bit
of a mystery. PMB
94
with a purplish-red spot or stripes in the middle of the

midlobe.
Encyclia cochleata: Regularly encountered in

the pond apple swamps, where it blooms mostly in the
fall and winter. Flowers typical of the triandrous
Florida mutation, with narrow, twisting greenish-yellow
sepals and petals dangling beneath the non-resupinate,
yellow-striped purple clamshell lip. There has been lots
of taxonomic juggling with this species. It doesn't fit in
the genus Encyclia in the strictest sense. The latest
proposal (Higgins, 1997) is to move it to the unwieldy
and unpronounceable genus Prosthechea. But that genus,
too, seems a catchall grouping, so the genus Anacheilium
eventually may prevail for the clamshell encyclias.
Polystachya concreta: Fairly common in pond

apple swamps and occasionally found in hammocks and
cypress forests. The bloom peak seems to be fall, but
some plants flower in the summer. The tiny, non-
resupinate flowers are conical and yellowish-green. This
species, now considered pantropical in distribution,
produces multiple secondary flower clusters along the
rachis of the inflorescence, accounting for the generic
name, which means "many spikes."
Harrisella porrecta: One of Florida's three

African-angraecoid-related leafless orchid species and
one of South Florida's two smallest orchids (the other
being the rare Lepanthopsis melanantha from the
Fakahatchee). A common small-twig epiphyte most
often seen on cypress trees, but also growing on a
95
number of other hosts, ranging from oaks to hog plum,

Ximenia americana, to old, woody stems of the large-
flowered climbing or swamp aster. Aster carolinianus. A
few tiny gray-green roots radiate from a short central
"stem," from which the thread-like inflorescences are
borne, mainly from late summer well into autumn. The
tiny yellowish flowers with their saccate nectar spur
appear quite complex under sufficient magnification.
The easiest way to search for this diminutive species is
to stand under a likely host tree and hunt for orchid-
looking roots. But plants are much easier to find in
fruit rather than in flower. The little globose green
capsules split open at the end to dehisce their seeds.
The walls of the capsules turn an orangy brown as they
mature, making the old capsules the prettiest part of the
plant to the naked eye. Because of the shape of the
seed capsules, this little orchid recently has acquired the
"common name" jingle bell orchid. At least it's a
common name that makes sense (compared to many
listed in Wunderlin, 1998). Some authors (Wunderlin,
1998) are using the name Harrisella filiformis for this
orchid, but Dr. James D. Ackerman in Puerto Rico, a
thorough botanist whose taxonomic judgments are to
be trusted when it comes to orchids, maintains that
Harrisella filiformis, is, in fact, a Campylocentrum species
different from the monotypic Harrisella porrecta
(Ackerman, 1995).
Polyrrhiza lindenii: Florida's legendary ghost

orchid. Another of our leafless African angraecoid
relatives. Surprisingly frequent in the pond apple
swamps around Rabenau Camp, in contrast to its
96
reported rarity elsewhere in and around the Big Cypress.

This species is amazing for the relatively large size of
the flower produced from such minimal plant mass.
The plant consists of a spidery array of fat, gray-green
roots with light green growing tips. These roots radiate
outward from the short central stem, which produces
the wiry inflorescence. In the Rabenau Camp region,
this species generally blooms from late June into
September. Because of the large white flower with
nocturnal fragrance and a very long nectar spur, the
pollinator is assumed to be a large, long-tongued night-
flying sphinx moth. Many authors are now using the
name Polyradicion lindenii for this species (Wunderlin,
1998.) Both names refer to the numerous roots often
produced by these orchids.
Ionopsis utricularioides: The only Oncidium

relative found at Rabenau Camp, where it is not very
common. These little twig epiphytes are most closely
related to the West Indian "equitant oncidiums" now
rightly segregated as the genus Tolumnia. The little
plants, sometimes reddish in intense sunlight, are found
most often on cypress trees, sometimes dangling off
their host by a few thread-like roots. The panicles of
pretty little white, purple-striped flowers generally are
produced from February through April.
Cyrtopodium punctatum: This is the most

spectacular orchid of Rabenau Camp, considering plant
size and the number of showy flowers produced. It is
found only sparingly in this area. It once may have
been more frequent in the region, but because of its
97
large size and showy bloom displays, collectors have

targeted it for years. Most of the plants seen around
Rabenau Camp have been growing low on cypress trees
in fairly sunny areas. It blooms primarily from February
through April. The flowers, to two inches across, are
crowded on panicles with large yellowish-green bracts
barred with reddish brown that mimic the sepals of the
flowers. The petals are yellowish with a scattering of a
few rust-red spots (accounting for the species epithet,
which refers to puncture wounds). The rather waxy lip
is a beautiful reddish mahogany color with a yellow
crest and center to the midlobe.
THE TERRESTRIALS
Spiranthes vernalis: The most common ladies'-

tresses orchid at Rabenau Camp, where it is at its bloom
peak in February and March. Flowering plants are often
found by the dozens on roadsides and in old pastures
and open prairies.
Spiranthes brevilabris: One small colony of

this rare species (rare for South Florida, at any rate)
persisted for a few years in an area of rough pasture
converted from pineland. First found on March 27,
1981, it was blooming at the same time as Spiranthes
vernalis, but the flowers seemed different, being smaller,
more yellowish and with an obviously shorter lip. The
plant, too, was quite different. Whereas Spiranthes
vernalis produces ascending, narrow, rather grass-like
leaves, those of this orchid were broad, rounded, and
98
formed a basal rosette, although they were starting to

fade at anthesis.
Using Dr. Carlyle A. Luer's distribution maps in

The Native Orchids of Florida (1972) as a guide, I deduced
that this orchid must be what he called Spiranthes
brevilabris var. floridana. But the flower description didn't
quite match up. That variety (now called Spiranthes
floridana) is essentially glabrous. But the flowers and
inflorescence of the plants near Rabenau Camp were
hairy, as clearly showed in the few rather poor-quality
close-up color slides I took on February 19, 1982,
before I "lost" this colony. Clearly, this must be
Spiranthes brevilabris var. brevilabris instead -- and the
Rabenau Camp-area population may be a first record
for Collier County, Florida3.
The only problem: Since I realized what this

orchid really is and sought to take better photographs
and collect a voucher specimen, I have been unable to
find it! I know the precise location, so that is not the
problem. The colony just seems to have "disappeared."
I'm hoping that, like isotrias in the North, the plants are
simply "taking a rest" (of many years duration!). But the
area were these plants occurred is becoming overgrown,
especially so since cattle were excluded from the
property. I'm wondering whether, if this pineland
burned (as is a common occurrence with this sort of
habitat), perhaps this colony might make a
2. This indeed is the first record for Collier County. PMB
99
reappearance. Nevertheless, year after year, the search

goes on ....
Spiranthes laciniata: This species is a near twin

of Spiranthes vernalis, with only minor floral details
separating the two. But what does separate them in the
vicinity of Rabenau Camp is a marked hiatus of about
two months between their bloom times, with Spiranthes
laciniata starting to flower in late May and extending into
July.
Flowering as they do at the start of the rainy

season in South Florida, the plants of Spiranthes laciniata
often bloom out of a few inches of standing water.
This species is common in the open glades of the
region, often becoming abundant along the edges where
the glades grade into open cypress forests.
Spiranthes torta: First found in flower on

June 23, 1996, this is the 28th orchid species discovered
in the Rabenau Camp region. It was blooming at the
same time as Spiranthes laciniata, which grew very nearby
in the same rough pasture succeeding to pineland. But
the center of the lip on flowers of this plant was
greenish, matching the plants of Spiranthes torta as I
knew it from verified populations on Big Pine Key in
the rock pinelands of the Lower Florida Keys. By
contrast, the lip of Spiranthes laciniata is yellowish in the
middle. Also, the flowers of this greenish-lipped plant
near Rabenau Camp were smaller, somewhat nodding,
with the lower flowers somewhat separated from the
others in the loosely spiraled raceme. The leaves were
100
grass-like, similar to Spiranthes laciniata, and not like Paul

Martin Brown's recently described Spiranthes eatonii
(Brown, 1999). A specimen was collected and
deposited in the herbarium at the Marie Selby Botanical
Gardens in Sarasota, Florida. But there is still some
small degree of doubt in my mind whether this is truly
S. torta. I will look for these plants in flower again this
year in the hope of alleviating these lingering doubts.
Spiranthes odorata: The only fall-blooming

ladies'-tresses we have found at Rabenau Camp, this one
is the whopper of the genus. Plants can grow to nearly
four feet tall. Luer (1972) treated it as a robust southern
variant of the northern nodding ladies'-tresses,
Spiranthes cernua, but it is now pretty well accepted as a
valid species in its own right. The tall stems of large
(well, large for a Spiranthes) white flowers occur most
frequently in open cypress forests, often growing out of
several inches of standing water, making this one of
Florida's "aquatic" orchids. The flowers, as the species
name implies, are quite fragrant, exuding a sweet, citrus-
like aroma during the day.
Stenorrhynchos lanceolatum: The only

segregate that has shown up at Rabenau Camp, this
species appears to be rare in that area. I have seen it
only once, and a plant collected by William Rabenau at a
second, widely separated station, serves as the voucher
for this species for area. This species is much more
abundant, especially along roadsides, slightly farther
north in peninsular Florida. Sometimes called the terra-
cotta orchid, that "common name" does not do justice
101
to the brilliant red-orange color of this species' large,

showy flowers (although, of course, color variations
occur, with even a green-flowered form). Both the
voucher specimen and the plant I photographed on Bill
Rabenau's western 50 acres were in bloom in late
March. Dr. Leslie A. Garay (1980), in his rather fine
splitting of the red-flowered Spiranthes, called this orchid
Sacoila lanceolata, and most authors now seem to be
following that usage.
Ponthieva racemosa: This intriguing little
spiranthoid orchid with the mysterious common name
shadow witch grows on a few tree islands, often near
the edge, where the island grades down into the
surrounding swamp. Perhaps they prefer this location
because of the greater abundance of moisture or
sunlight -- or both. However, the first plants of this
species recorded for the Rabenau Camp area grew
under cabbage palms (Sabal palmetto) at the edge of a
pineland. But, again, it was a fairly moist habitat. I
recently returned to this spot after many years, only to
find the whole understory rooted up by feral hogs.
Plants of this species produce basal rosettes of beautiful
velvety green leaves. They always remind me of the
northern goodyeras - minus the beautiful tracery of
veins in the leaves. In the Rabenau Camp region, this
delightful little species produces its intricate white,
green-striped, upward-facing flowers primarily in
January
and February.
102
Zeuxine strateumatica:: This little naturalized

exotic orchid "weed" is found at Rabenau Camp mostly
in disturbed areas, such as roadsides and old pastures.
It is native to southeastern Asia and is thought to have
come into Florida with seeds of centipede grass,
Eremochloa ophiuroides, imported in the early part of this
century for use as a grass for lawns (hence, the rather
prosaic common name lawn orchid applied to the
Zeuxine). The orchid apparently liked its new home
because it is now reported from most of Florida's 67
counties, as well as from areas of a few adjacent states.
The little pointed clusters of glistening white flowers
with their golden yellow, spade-shaped lips are quite
pretty under magnification. They are borne mostly in
the winter months.
Oeceoclades maculata: Called Eulophidium

maculatum when it was first detected in Florida in the
early 1970s, this recent arrival is now so happy that is
growing like a weed in the southern part of the state.
Unfortunately, that proliferation has gotten it into
trouble, earning this species the label "exotic pest plant"
from a quasi-governmental group called the Exotic Pest
Plant Council. Because the group has put this orchid
on their hit list of "bad" plants, well-meaning but
misguided "environmentalists" are ripping this orchid
out of the ground wherever they find it and destroying
the plants. The laugh is on them, though, because
whether this orchid got here by natural means or
through man's intervention (a point still much debated),
Oeceoclades maculata is firmly entrenched in our state's
103
flora. It is here to stay, and these environmental "do-

gooders" will never eradicate it.
This orchid of African origin is the 29th and

latest species recorded in the vicinity of Rabenau Camp.
A few young plants were discovered in June of 1997 on
a tree island that a group of us have to cross to get to a
deep swamp harboring a healthy population of ghost
orchids, and I wonder whether we inadvertently may
have spread the seeds of Oeceoclades there on our muddy
boots from other sites where this species is plentiful.
No sexually mature plants of Oeceoclades have turned up
on this island yet, but it's only a matter of time until we
find plants in flower.
Elsewhere in South Florida, this species blooms

from late summer into autumn. This is the last part of
the rainy season, and it is theorized (Gonzalez-Diaz and
Ackerman, 1988) that the weight of raindrops hitting
the flowers jars the pollinia loose and onto the flower's
own stigma. Our plants of this species, then, would be
self-pollinating, thus accounting for the heavy fruit set
and resulting rapid territorial expansion of this orchid.
Eulophia alta: Related to Oeceoclades maculata

and, like it, a plant of African origin that is now
widespread in the Western Hemisphere, this species is
fairly common at Rabenau Camp, growing in disturbed
areas, such as roadsides, was well as in open natural
habitats. The sterile plants resemble palm seedlings,
accounting for the common name wild coco (as in
coconut).
104
This fall-blooming species is one of our showiest

native terrestrial orchids, producing inflorescences to six
feet tall with dozens of flowers. The species epithet, in
fact, means "tall." The three sepals, distinctively
pointing upward, range in color from greenish to honey
gold to mahogany. The lateral petals are usually
similarly colored and converge with the crested lip to
form a tunnel, presumably for pollinator alignment.
The lip can be purplish to pure white in color (the latter
color form being extremely rare).
Triphora gentianoides: Aesthetically one of the

least appealing orchids of the Rabenau Camp area, this
species was first discovered there in early July of 1984
on the same tree island where we later found Oeceoclades
maculata. Sizable Triphora colonies persist on this island,
and, in the past, a small colony was found on a nearby
island, but that spot hasn't been revisited in a number of
years. Although this species is frequent in yards and the
mulch around landscape plantings at spots along the
southeast Florida coast, the 1984 collection of this
species on the tree island near Rabenau Camp may
represent the first record of Triphora gentianoides for
Collier County Florida4.
Malaxis spicata: This delightful miniature

produces the second smallest flowers of any orchid
species encountered in the vicinity of Rabenau Camp.
To call this species a miniature may be a little misleading
4
Also a first record for Collier County. PMB
105
because robust inflorescences can approach nearly a

foot in height at the end of a long period of blooming,
although most plants are considerably smaller. Plentiful
in the Rabenau Camp region, this species generally
grows in swamps under quasi-epiphytic conditions in
the accumulated detritus on
stumps, fallen logs, and the knees and buttressed bases
of cypress trees, often just above the high water line. It
blooms primarily in autumn. In the Rabenau Camp
area, it also has been found rooted in the sandy humus
under wax myrtles, Myrica cerifera, in areas that have only
a short hydroperiod.
The two leaves and inflorescence emerge from a

small cormous base. The inflorescence continues to
elongate as more and more of the tiny greenish to
bronzy orange flowers are produced over a long period
of time. The pointed lip is uppermost due to a 360-
degree twist. Dr. Robert L. Dressler (1981) calls the
flower "hyper-resupinate," and Luer (1972) theorizes
that this unusual occurrence may be due to adaptation
to a changing pollinator.
Habenaria quinqueseta: Four true habenarias

occur in Florida (or five if you support the separation of
Habenaria macroceratitis). Of the three that occur at
Rabenau Camp, Habenaria quinqueseta is arguably the
prettiest. There, it mostly grows in slightly higher, drier
areas of South Florida slash pine, Pinus elliottii var. densa,
where it generally is at its bloom peak from late August
into September.
106
The flowers, ranging from white to greenish

white, are typically shaped for most of the genus
Habenaria in the New World. Although the species
name means "five bristles," referring to what at first
appears to be a lip of five narrow, sweeping segments,
the labellum, in fact, is only three-lobed. The
uppermost segment on each side of the "five-lobed" lip
is actually the lower lobe of each deeply bilobed lateral
petal. The upper lobe of each lateral petal tucks under
the edge of the concave dorsal sepal to form a hood
over the entrance to the inch-long (or longer) nectar
spur growing from the back of the lip. Habenaria
quinqueseta is not common at Rabenau Camp, perhaps
due to the limited amount of pineland in the area.
Habenaria odontopetala: This is the most

common Habenaria species at the southern end of
Florida, where it generally blooms later than Habenaria
quinqueseta, with flowering starting in midautumn. Some
of the geenish-yellow flowers persist nearly through the
winter, although they don't appear to be reproductively
active by that time.)
The flowers, at first glance, appear to be

differently formed from those of Habenaria quinqueseta,
but on closer examination, the common Habenaria
flower pattern is evident. The typically long side lobes
on the lip of most habenarias here are reduced to nubs
at the top of the slightly broader midlobe. The upper
lobe of each lateral petal is broadly toothed at the apex
(accounting for the species epithet and the common
name, tooth-petaled rein orchid), and these lateral
107
petal lobes often are more erect, standing out somewhat

from the concave dorsal sepal. The lower lobe of each
lateral petal, like the side lobes of the lip, is reduced to a
tiny lobule, sometimes barely noticeable.
At Rabenau Camp, plants of Habenaria

odontopetala can be quite robust in sunny optimum
conditions, occasionally growing to four feet tall with
inflorescences densely packed with flowers. But in the
species' more common shady hammock-type habitats,
plants are much smaller, producing only scattered
flowers on the inflorescence. Wunderlin (1998) uses the
name Habenaria floribunda for this orchid.
Habenaria repens: This is the most common

Habenaria in Florida (and perhaps all of the Western
Hemisphere). Ironically, it is found only sparingly in
southern Florida. I have seen it only once in the
Rabenau Camp region, when I photographed a stem
from a single colony on November 1, 1980. I have
searched for it in that area ever since, but to no avail.
This species is, however, encountered occasionally in
the Fakahatchee Strand to the west.
The plants seen near Rabenau Camp grew among

pickerelweed, Pontederia cordata, in an open swampy area
at the edge of a tree island far to the south of camp.
That is a clue to the habitat of this species, the most
aquatic of Florida's native orchids. The plants often
grow and bloom in floating mats of vegetation in lakes
and roadside ditches. That habitat, coupled with the
myriad little green, typical Habenaria-shaped flowers,
108
accounts for the apt common name water spiders.

Elsewhere in the state, this species appears to bloom
sporadically throughout the year.
Platanthera nivea: This attractive and sweetly

fragrant little orchid is found sparingly in the region of
Rabenau Camp, where it grows in open glades, often
near the edges of cypress forests. All the plants we have
encountered have been well to the southwest of the
main camp, closer to typical Big Cypress habitat.
Called the snowy orchid for good reason (even

its species name alludes to this), the glistening little blue-
white flowers are borne in conical clusters at the tip of a
thin blue-green stem. The lip is uppermost in this
species, although this can be hard to detect at first
glance. The dorsal sepal, lowermost in these is slightly
broader and imitates the pollinator "landing pad"
function normally performed by the labellum. The
lateral sepals, located where you'd expect the lateral
petals to be, have an odd twist to them so that for most
of their length, you are looking at their reverse side.
The long nectar spur juts straight out horizontally
behind each slower. At the center of the flower is a
yellow spot on the column.
In the Big Cypress, this species seems to have a

rather restricted bloom period from late May into June.
Blooming at the outset of the rainy season, the plants
often grow out of a few inches of water, much like
Spiranthes laciniata, which flowers at the same time in
similar habitats.
109
Dr. Richard P. Wunderlin (1998) has, rather

inexplicably, returned this orchid to the genus Habenaria,
where it is clearly out of place. Even in Platanthera, it is
a bit of an oddball, so we eventually may see another
generic name for this pretty orchid.
Calopogon tuberosus: Like Platanthera nivea,

this orchid is found sparingly in the vicinity of Rabenau
Camp, usually well to the southwest of camp, where it
grows in open glades. It is well represented in similar
habitats in the heart of the Big Cypress. This is easily
one of our prettiest native terrestrial orchids and is a
shame it lost the apt species name pulchellus, which
means "pretty." This is probably the most widespread
orchid of eastern North America, growing from
Newfoundland south to Florida, Cuba and the
Bahamas, and from the Atlantic westward to beyond the
Mississippi. Because this orchid is so well known, it
does not need any further description here. Plants can
be found in flower in the Big Cypress anywhere from
March to June. The southern form in these prairies has
been called var. simpsonii by some authors.
Bletia purpurea: A common species at Rabenau

Camp, where it grows on canal spoils banks, along
roadsides, in old pastures, on floating logs in swamps,
and even on cypress knees and the buttressed bases of
cypress trees.
Blooming from winter into spring, it produces

tall, sometimes branched, inflorescences of pretty pink
110
flowers. When the flowers are open, the sepals are

arranged in a typical triangular orchid pattern. The
petals, however, arch over the lip, forming a tube,
probably for pollinator alignment, much as in Eulophia
alta in South Florida or Cleistes, Isotria or Pogonia species
elsewhere. The magenta-edged lip has beautiful raised
golden keels running down much of the length of the
midlobe. In some Florida mutations of this species, at
least one of these yellow keels is repeated in the lateral
sepals.
Because this tropical orchid is at the northern

end of its range in southern Florida and sufficient
suitable pollinators may not be available, many, many
plants of this species in Florida produce cleistogamous
flowers, that is, the buds mature and self-pollinate
without ever opening to reveal their beauty to the
world. What a waste, aesthetically. This shortcut to
colonization may be successful at first. But will the
long-term results be beneficial genetically?
Although he was a good steward of the land, Bill

Rabenau was forced to sell his camp property when the
Big Cypress National Preserve expanded its
northeastern borders. He bought new property in
adjacent Hendry County, west of the Big Cypress Indian
Reservation, and built a sumptuous new "camp" in a
shady hammock of live oaks, Quercus virginiana and
cabbage palms. But it was never the same to him. He
loved his old camp down in "the Cypress," as he called
it, and its loss through the forced buyout broke his
heart. He lived to enjoy his new camp only a short
111
while. Bill Rabenau died on April 16, 1991, and this

paper is dedicated to his memory.
After Bill's eviction from Rabenau Camp, I

despaired of ever having the pleasure of botanizing
there again because one of the first things government
bureaucracies do when they buy public land is deny the
public access to it -- even though it was bought with the
public's money. Luckily, that did not occur with the
property around Rabenau Camp, so in July of 1993, we
revisited the area after several years. Since then, I have
visited frequently. Of course, the camp structures are
gone. They were bulldozed into a pile and burned. But
the natural areas still gloriously intact, and I have been
able to add Spiranthes torta and Oeceoclades maculata to our
list of orchid species for the region since then.
Also, the Big Cypress National Preserve has been

kind enough to grant to me a limited collecting permit
so I can complete my herbarium voucher series for the
orchids of Rabenau Camp. Of the 29 species we have
discovered there over the past 20 years, I still need
herbarium specimens for just six: Epidendrum difforme,
Epidendrum nocturnum, Habenaria repens, Harrisella porrecta,
Oeceoclades maculata and Spiranthes brevilabris. I hope to
add some of these this fall. For the moment, though, I
fear I'll never get my voucher specimens of Habenaria
repens and Spiranthes brevilabris, for reasons discussed
earlier in this article. All voucher specimens are on
deposit in the herbarium at the Marie Selby Botanical
Gardens in Sarasota, Florida.
112
And who knows? Perhaps as I continue to

search for these "missing" voucher specimens, I may
encounter still more orchid species for Rabenau Camp.
FOR FURTHER READING:
Ackerman, James D. 1995. An Orchid Flora of Puerto Rico and the
Islands. The New York Botanical Garden, Bronx, New
York.
Austin, Daniel F. (undated). Vascular Plants of Fakahatchee Strand
State Preserve. Naples Chapter, Florida Native Plant
Society.
Brown, Paul Martin. 1999. Spiranthes eatonii Ames ex P.M. Brown:
Eaton's Ladies'-Tresses, North American Native Orchid
Journal 5(1):3-15.
Dressler, Robert L. 1981. The Orchids: Natural History and
Classification. Harvard University Press, Cambridge,
Massachusetts.
Garay, Leslie A. 1980. A Generic Revision of the Spiranthinae,
Botanical Museum Leaflets (Harvard University), 28(4): 277-
425
Gonzalez-Diaz, Nancy, and James D. Ackerman. 1988.
"Pollination, Fruit Set, and Seed Production in the Orchid
Oeceoclades maculata, Lindleyana 3(3):150-155.
Hagsater, Eric. 1993. Epidendrum floridense Hagsater, Icones
Orchidacearum 2, Plate 133.
Higgins, Wesley E. 1997. A Reconsideration of the
Genus.Prosthechea (Orchidaceae). Phytologia, 82(5):370-383.
Luer, Carlyle A. 1972. The Native Orchids of Florida. The New
York Botanical Garden, Bronx, New York.
McCartney, Chuck. 1981. The Orchids of Rabenau Camp: 1. The
Epiphytes. American Orchid Society Bulletin 50(5):527-535.
1981. The Orchids of Rabenau Camp: 2. The Terrestrials.
American Orchid Society Bulletin.50(6):653-660.
1986. More Orchids at Rabenau Camp. American Orchid Society
Bulletin, 55(3):232-241.
Wunderlin, Richard P. 1998. Guide to the Vascular Plants of Florida.
University of Florida Press, Gainesville.
113
Chuck McCartney, 2226 Lincoln St., #3, Hollywood FL 33020

Chuck is a prolific writer and explorer of the orchids of the
Southeastern United States. He is a former editor of the
American Orchid Society Bulletin and a popular speaker and field
trip leader with the various natural history organization
throughout Florida.
114
Sheviak: PLATANTHERA HYPERBOREA AND A REAPPRAISAL OF
GREEN PLATANTHERAS
PLATANTHERA HYPERBOREA AND A

REAPPRAISAL OF GREEN
PLATANTHERAS
Charles J. Sheviak
The American members of Platanthera that were

assigned to the genus Limnorchis by Rydberg (1901)
constitute some of the most baffling taxonomic subjects
in our flora. Portions of the group seem to defy
reasonable delimitation of species, and it is indeed
possible that no truly acceptable species concept will be
found to be applicable to them. Foremost among the
problematical plants are those comprising the so-called
Platanthera hyperborea - P. dilatata complex. I've been
studying these plants rather intensively for 20 years, and
in a more general way for over 15 before that. I've
studied populations across most of the range of the
complex from northwest Alaska to the Mexican border
to the Atlantic Coast. Representative specimens have
been maintained in cultivation for periods ranging to
over 15 years. Cytology and pollination have been
investigated. A synthesis of the information gained has
resulted in some progress, and a series of publications
on the group is now beginning to appear. The present
115
GREEN
McCartney: PLATANTHERAS
paper presents an overview of certain patterns of

variation that have become apparent.
For 25 years now we've been using Carlyle Luer's

treatment (Luer, 1975) of the group as the basic
reference for the complex. I t has indeed been very
helpful, and Luer brought attention to various taxa that
had been all but lost in Correll's earlier treatment that
included fewer, more broadly delimited taxa (Correll,
1950). Now, with considerably more information and
experience, further revisions are warranted. Luer's
framework is still a sound foundation upon which to
build, however, and it will be used here as an outline for
the present paper.
I want to introduce the group with its most

unmistakable member, Platanthera dilatata (Pursh) Lindl.
ex Beck. This species is distinctive in its brilliantly pure
white flowers; in all other species they are more or less
green. Platanthera dilatata furthermore has a strong clove
scent that is characteristic and apparently limited to this
species. The lip typically is marked by a broad, rounded
basal dilation, although the degree to which it is
expressed varies; most commonly it is suborbicular to
transversely elliptical. The species is transcontinental in
the north and widespread in the western cordillera. Like
much of the rest of the group, it is quite variable, and
traditionally three varieties have been recognized.
116
Sheviak: PLATANTHERA
Sheviak: PLATANTHERA HYPERBOREA
HYPERBOREAAND
AND
A REAPPRAISAL
A REAPPRAISAL
OFOF
Sheviak: PLATANTHERA HYPERBOREA
GREEN HYPERBOREA
GREENHYPERBOREA
AND
AND
PLATANTHERASAND
A REAPPRAISAL
PLATANTHERAS A REAPPRAISAL
A REAPPRAISAL
OFOFOF
GREEN
GREEN
GREEN
PLATANTHERAS
PLATANTHERAS
PLATANTHERAS
In the boreal forest east of the cordillera, ranging

south through the Great Lakes Region and eastward to
the Coast, is the nominate Platanthera dilatata var. dilatata.
This variety is distinguished from other varieties by a
slender, mostly white spur about the length of the lip.
The viscidia are linear.
From southern Alaska southward through the

Rocky Mountains to Colorado, we find Platanthera
dilatata var. albiflora (Cham.) Ledeb. This variety is
characterized by a short, clavate, generally green spur.
The viscidia are a bit broader than in var. dilatata, linear-
oblong to oblong. The lip is also often broader than in
the nominate race, and in Alaska the basal dilation is
sometimes so broadly elaborated that the base of the lip
actually bears two forward-directed lobules.
To the west of the Rocky Mountains is Platanthera

dilatata var. leucostachys (Lindl.) Luer. This variety is
distinguished by a slender white spur markedly longer
than the lip. Often the lip is very slender, sometimes
narrowly lanceolate and undilated, and the viscidia are
linear. Most significantly, in extreme form, such as in
the Sierra Nevada, var. leucostachys is scentless during the
day and fragrant at night.
Platanthera dilatata var. leucostachys is obviously

specialized for moth pollination, whereas the short
spurred, diurnally fragrant var. albiflora seems to be
117
McCartney: GREEN PLATANTHERAS
specialized for diurnal, shorter-tongued insects. The

short, fat spurs of the latter variety reach their greatest
reduction in the southern Rockies of Colorado, where
they are sometimes reduced to a small green sac.
Northward they are less reduced. This is a recurring
theme in the group: local specialization and variation.
Although in extreme form Platanthera dilatata var.

leucostachys is far-western, plants referable to it range
eastward across the Great Basin to western Utah. I say
'referable', because these more eastern plants often don't
exhibit the pronounced nocturnal fragrance, it being
more variably and weakly expressed. Indeed, eastward
from the Sierra the plants become less distinctive. For
instance, the Ruby Range in northeastern Nevada
supports the species in moist subalpine meadows and
along streams at lower elevations. Spur length in these
populations is variable, and many plants would
arbitrarily be assigned on this basis to P. dilatata var.
ideally, whereas others are referable to var. leucostachys.
Varieties leucostachys and albiflora therefore appear to be
geographically variable, reaching extreme form in
restricted geographic areas. In fact, northward the
situation becomes quite complex.
In interior British Columbia most plants bear

spurs that are short, stout, and green when the flowers
open. In cultivation, however, as flowers age the spurs
commonly elongate, becoming slender, white, and
about equaling the length of the lip. Variety albiflora was
118
GREEN
described from Unalaska, and indeed, most Alaskan

plants seem referable to this variety, with short, fat,
green spurs. A few herbarium specimens show spurs
suggesting var. dilatata, but it's not clear that that is what
they are. I've grown a few plants of variety albiflora from
the Kenai Peninsula. These were all representative var.
albiflora when collected, and that is what they looked like
in subsequent years. One year, however, one plant
opened its characteristic var. albiflora flowers, and then
after an interval of a few days, the spurs abruptly
elongated until they were slender and markedly longer
than the lip. Essentially the plant changed from var.
albiflora to var. leucostachys. How then do we interpret the
long-spurred plants from the Alaskan Panhandle, where
var. leucostachys seems to reach its northern range limit?
Basically, it seems that the Northwest is the center

of variability of Platanthera dilatata, and as the species
ranges eastward through the boreal forest, and
southward down the Rockies and the more western
ranges, variability is reduced in response to
specialization for different pollinators. It seems then
that the recognized varieties of P. dilatata are really just
endpoints in a very complex variation pattern. They
have some utility for discussion purposes, but they are
very simplistic representations of the underlying
situation.
The very distinct Platanthera dilatata is therefore
119
GREEN
quite complex, perhaps unexpectedly so. The rest of the

members of the complex are the green-flowered plants
that I refer to as "basic green platantheras," or "BGP's"
for short. These are the plants that are so often
troublesome, although some are actually very distinctive
and readily determined once they, and their
characteristics, are recognized.
Platanthera hyperborea (L.) Lindl. has the most

extensive range of any species in the group, ranging
from northwestern Alaska southward to New Mexico
and southeastward to central Illinois and New England.
Although this appears to be a very nondescript plant, it
is actually one of the most distinctive members of the
group. The markedly yellowish lip is rhombic-lanceolate
in outline, not at all dilated. The spur is decidedly
clavate and slightly shorter than the lip, but it appears
shorter because of a sharp curve. The column of this
species is very distinctive. It is proportionately very
small, with the anther sacs lying at low angles atop the
stigma. Most remarkably, the flowers are self-
pollinating: the pollinia fall free of the anther sacs and
eventually contact the stigma. Perhaps because of this,
across most of its range the flowers are scentless. The
pollinaria are functional, however, and the viscidia are
orbicular. None of our other recognized species have
such a column nor pollination mechanism. The
combination of column structure, pollination
mechanism, lip color and lip and spur shape, and usually
scentless flowers combine to make P. hyperborea very
distinctive.
120
GREEN PLATANTHERAS
It is ironic, then, that two of the most distinctive

and well-defined members of the group have been used
to designate the so-called hyperborea-dilatata complex.
The complex is actually composed of other species and
plants.
Luer's treatment recognized five varieties of

Platanthera hyperborea, and they form a useful framework
for presentation of some of the remaining plants of
interest here. Variety hyperborea has just been introduced.
Platanthera huronensis (Nutt.) Lindl. has been one of

the most vaguely-defined concepts in the group. Luer
treated it as a variety of P. hyperborea, in keeping with
most earlier workers who recognized it. This plant
immediately differs from P. hyperborea in its whitish
green flowers. The lip is generally rounded-dilated at the
base, but generally less so than is characteristic of P.
dilatata. The spur is slender and about the length of the
lip. The flowers are intensely sweetly pungent-scented
with a scent that to me suggests a sweet version of the
laundry detergent isle at the grocery store. The column
is larger and the anther sacs much higher than in P.
hyperborea; the flowers are not self-pollinating. The
viscidia are oblong.
121
GREEN PLATANTHERAS
This species has been the source of a lot of

confusion, stemming in part from its color and the
dilation at the base of the lip. It looks like a hybrid of
Platanthera dilatata and P. hyperborea, and very likely that's
how it arose. The species is a tetraploid, and tetraploid
species in nature are almost always of hybrid origin.
Very likely, then, P. huronensis arose as a P. dilatata
hybrid, and Catling and Catling (1997) presented results
of a numerical analysis indicating that P. hyperborea might
be the other parent.
The species ranges southward in the cordillera

through British Columbia to Colorado and Utah, and
across the Great Lakes region to the Atlantic Coast. The
plants vary greatly in stature and can be quite robust,
reaching over a meter in height. The density of the
inflorescence varies from lax to very dense.
Two other points can be made here. First, in the

northeast this species is commonly confused with
putative hybrids of Platanthera dilatata and P. hyperborea
and called P. xmedia (Rydb.) Luer. Secondly, the plants
in western Canada and Alaska that Luer called P.
hyperborea var. viridiflora (Cham.) Luer are in all respects
referable to P. huronensis.
The next species I want to discuss is Platanthera

purpurascens (Rydb.) Sheviak & Jennings. This is the
fourth taxon treated by Luer as a variety of P. hyperborea.
Rydberg's name had been ignored for most of this
century, and we owe Luer a lot for introducing the
122
GREEN
plant, which had been quite problematical for many

years. Again, as BGP's go, P. purpurascens is quite easy to
recognize once you learn what to look for. The
vegetative habit is distinctive, with rather short, blunt,
abruptly wide-spreading leaves. This spreading leaf habit
is a significant feature; it doesn't seem to vary with the
amount of exposure to sun and wind. The flowers tend
to cluster in fascicles in a long, slender raceme, a
particularly unusual and significant feature. Lips vary in
shape from narrowly lanceolate to broadly dilated, and
in color from watery greenish yellow to intensely bluish
green. Sometimes reddish blotches occur on the lip or
other floral segments; these are contributed by the
coloring of dense networks of minute veins and
evidently are responsible for the specific epithet.
Regardless of the actual color, the intense, watery
quality is characteristic. The spur is variously short and
inflated, usually strongly inflated-clavate to saccate and
scrotiform. The flowers are strongly and uniquely
scented with a marked musty, semen-like scent.
Columns are rather typical of the group, with the rather
small anther sacs well-elevated above the stigma and the
rostellum lobes somewhat spreading. The viscidia are
typically orbicular, but vary to oblong. There is quite a
bit of variability in these features, but the uniformly
short spur and peculiar coloring, abruptly spreading
leaves, and singular scent hold the group together and
indeed make it quite obvious.
123
GREEN
I really don't think Platanthera purpurascens has

much to do with P. hyperborea, but it shares a few
features with P. stricta Lindl. Platanthera stricta is a more
northern species, ranging from Alaska southward to
northernmost California and about Yellowstone
National Park. Throughout this range the plant is
marked by short, abruptly spreading leaves. The spur is
relatively short and inflated, and varies from inflated-
clavate to very short and saccate. Lips are generally
rather long and linear to merely lanceolate. The color,
however, is a flat green, not the watery, deep coloration
of P. purpurascens, and the column is very different than
in that species, with prominent, more or less parallel
anther sacs. The flowers are, to my nose, virtually
scentless, sometimes with just the slightest spicy quality.
Viscidia are orbicular.
Taxa in this group are obviously capable of

considerable variation, regardless of how strictly they
might be delimited, and interpretation of limits is
confounded by convergence under similar pollination
pressures. The similar range of spur forms in Platanthera
purpurascens and P. stricta is consequently of uncertain
significance. Their vegetative habits might be more
significant, but can a single vegetative feature define a
species? I think that the very different columns of P.
purpurascens and P. strict a, in contrast, lie outside the
range of variation seen in other species in the group
and, when taken together with the other differences
between them, indicate that, indeed, we are dealing with
two distinct species.
124
GREEN
That said, I might point out yet another problem.

From southern British Columbia to northern California,
at least, occur very tall, slender plants with less abruptly
spreading leaves. The spurs on these plants are
uniformly inflated-clavate, not at all saccate.
They are similar to those often seen on

uppermost portions of the inflorescences of the more
typical Platanthera strict a discussed above, so they are not
unique. However, in addition to these morphological
differences, these slender plants have proven to be
exceedingly difficult to cultivate under my conditions,
whereas the more typical form grows reasonably well
for me. Hence, these two groups may be different at
some level.
That brings us to Luer's Platanthera hyperborea var.

gracilis (Lindl.) Luer, the last of his varieties of this
species. He illustrated this variety with a single photo
from Yellowstone. Comparison of Luer's photo with
Lindley's type at Kew, collected near the coast of British
Columbia on the border with the end of the Alaskan
panhandle, discloses very different plants. The holotype
sheet includes three specimens, all with very narrow
leaves that in one specimen are quite long. The plants
otherwise seem to fall nicely into P. stricta. The leaves
evidently have a tendency to spread abruptly. Lips are
long and slender and the spurs are entirely comparable
to those of the slender, problematical plants of P. stricta
mentioned above. I think P. gracilis Lindl. is refereable to
P. stricta, at least at some level.
125
McCartney: ORCHIDS
Sheviak: PLATANTHERA AT RABENAU
HYPERBOREA CAMP
AND A REAPPRAISAL OF
GREEN PLATANTHERAS
This may dispense with the name, but what was

Luer's plant? His published photo shows a slender
inflorescence, lanceolate lips, and blunt, clavate spurs a
bit shorter than the lip. I have found entirely
comparable plants in northern Colorado, geographically
not too far removed from Luer's locality. From what
can be seen in Luer's photo, the match between the lip
and spur of these plants is precise. These plants also
bear 'normal' columns: they are not merely Platanthera
hyperborea with slender inflorescences.
After years of looking for, and at, these things in

the field, I've become convinced that the plants
represented by Luer's photo are products of
hybridization of Platanthera dilatata var. albiflora and P.
purpurascens. The plants designated gracilis Luer
commonly occur in complex populations with these two
species and often with a variety of other peculiar plants
that exhibit considerable variation in lip shape and in
flower color, varying from deep green to dull white.
They are uniformly fragrant, usually with the
pronounced sweet-pungent scent of P. huronensis,
sometimes a bit more musty as in P. purpurascens.
Flowers often tend to be grouped in fascicles. Viscidia
are oblong. Individual plants furthermore may vary
greatly from year to year. One plant that I maintained in
cultivation for some years was essentially identical to the
plant in Luer's photo when collected and subsequently
126
GREEN
in cultivation, except for one year when it produced a

very dense inflorescence and virtually duplicated P.
huronensis in all salient characters. Unlike that species,
however, 'this plant, like the others discussed here, was
diploid. These plants quite obviously constitute hybrids
of the two diploid species with which they commonly
grow, and between which they vary.
To date I have found only a single specimen that

was not diploid, and this was triploid, as reported
previously (Sheviak & Bracht 1998). This plant, growing
with various diploid intermediates and the parental
species, suggested Platanthera huronensis and differed
from that species primarily in a very short spur and an
abruptly dilated lip with an eroded margin. The
morphology and cytology of this plant suggest a hybrid
of P. huronensis and P. dilatata var. albiflora, but the
former was not evident at the site. Of course, it may
have been present but vegetative, or it may have
occurred there in the past. Indeed, it would have taken
only a pollen grain brought in from another colony to
have given rise to this plant.
Demography, then, is a complicating factor that

contributes to the difficulty of interpreting populations.
The plants blooming one season do not necessarily
represent the total set of variants present then or in the
past. Indeed, by revisiting populations over periods of
up to fifteen years, I've noted marked changes in
127
abundance of Platanthera dilatata, P. purpurascens, and the

evident hybrids. In some years, colonies which normally
supported a complete range of parentals and
intermediates were wholly lacking in blooming plants of
one of the parents or the hybrids. Without such a
temporal sample, the total variation pattern and it's
significance can easily be missed.
The evidence suggests that the plants identified by

Luer as Platanthera hyperborea var. gracilis are products of
hybridization. If instead they are some distinct entity of
their own, they are so thoroughly involved in
hybridization with other species that it is difficult to
justify maintaining them as distinct. With that in mind,
the situation on Mt. Graham in southeastern Arizona,
where P. purpurascens occurs at perhaps its
southwestemmost station, may be significant. Also on
Mt. Graham are plants with the same morphology and
sweet scent as the Colorado plants discussed above. If
these things are hybrids, what are they hybrids of?
There isn't another species on Mt. Graham, nor

anywhere remotely near it. The nearest P. dilatata is in
northernmost New Mexico. These Arizona plants might
suggest that Luer's plants are indeed a distinct species
(or whatever), and in portions of their range they occur
with other species, perhaps hybridizing with them or at
any rate being confused with hybrids. On the other
hand, two other explanations are equally plausible. First,
these southern populations could represent the effects
of gene flow in the past, when another species was
present.
128
GREEN
Certainly the Pleistocene provides a ready

explanation: P. dilatata almost certainly occurred in the
area under the cooler, more moist climate that prevailed
in the Southwest at that time. Or, secondly, perhaps P.
purpurascens simply varies more than I've realized.
Perhaps these indeterminable BGP's, or some of them,
represent merely extreme forms of P. purpurascens.
A population of similar plants in the central Sierra

Nevada Mountains provides further ambiguous but
provocative evidence. I was directed to this site by
Ronald Coleman, who, following Luer, referred the
plants growing there to Platanthera hyperborea var. gracilis.
Throughout the area P. dilatata var. leucostachys was a
common plant, but in only one small seep occurred
green-flowered plants with lanceolate lips, oblong
viscidia, and slenderly clavate spurs about the length of
the lip. The plants appeared very similar to the plants
discussed above, except that the spurs were slightly
longer and the flowers wholly scentless. Subsequently in
cultivation, the lips were longer and proportionately
more slender, and the spurs proportionately longer and
more pointed. Lip color, too, changed from a medium,
flat green to a watery yellowish green.
What are these plants? This colony is not unique,

for I have seen similar herbarium material from a few
other sites in the region. At the station I visited, the
indeterminable green-flowered plants seemed to be
hybridizing with Platanthera dilatata var. leucostachys,
producing greenish white flowered plants with longer
spurs. Platanthera purpurascens is known from a few
129
disjunct populations in the same area; its occurrence

there may be significant, or it may be coincidental.
Despite the ambiguity, the distribution pattern of these
indeterminable BGP's suggests a plausible origin.
An hypothesis that accommodates all of the

problematical plants resembling the plant figured by
Luer as Platanthera hyperborea var. gracilis rests on one
question: what is the most generalized flower in this
group, one without any particular specializations? It is
medium green with lanceolate lip, clavate spur a bit
shorter than the lip, and probably oblong viscidia.
Deviation from this norm in any of these features, as
characterizes the various species in the complex, thus
represents a derived condition. For the most part this
probably is due to specialization for a particular type of
pollinator, but in some cases it might result merely from
drift. Anything that disrupts the genetic balance that
underlies the specializations characterizing the various
species is apt to produce a nondescript, basic flower,
like the one figured by Luer. This can be hybridization,
because genetic recombination may simply erase derived
states based on homozygous conditions.
Furthermore, the physiographic complexity of

the cordilleran region provides a mechanism for
generating considerable variability within species. More
generally, then, as species reach their range limits or
occupy habitats outside the norm, the selective
pressures that maintain their distinctive features may be
lessened or absent, and populations then may drift back
to a basic floral form. Hence, across the region,
130
GREEN
different processes acting on different species may have

given rise to very similar plants.
Considerable progress has been made in our

understanding of the so-called Platanthera hyperborea-
dilatata complex, and this gives hope that we may
ultimately arrive at a satisfactory taxonomy.
Nonetheless, the remaining known problems are
daunting, and each year in the field brings forth new
ones. Whatever the practical taxonomic products
ultimately prove to be, however, the very complexity of
this fascinating group suggests its significance as an
evolutionary research tool. In essence, these
problematical BGP's may be convergence products that
have arisen due to an abatement of selective pressures
rather than through their common action.
Literature Cited
Catling, P.M. and V.R. Catling. 1997. Morphological discrimination of
Platanthera huronensis in the Canadian Rocky Mountains. I.indleyana 12: 72-78.
Correll, D. S. 1950. Native Orchids of North America North of
Mexico. Chronica Botanica.
Luer, C. A. 1975. Native Orchids of the United States and Canada, excluding Florida.
New York Botanical Garden.
Rydberg, P. A. 1901. The American species of Limnorchis and
Piperia, north of Mexico. Bull. Tore.. Boot. CI. 28: 605-643.
Sheviak, C. J. & M. Bracht. 1998. New chromosome number
determinations in Platanthera. No. Am. Native Orchid
Journ. 4:168-172.
Contribution number 794 of the New York State Museum.
Charles J. Sheviak, Biological Survey, New York State Museum,
Albany, NY 12230. Chuck is one of the leading authorities in the native orchid
community and is has published extensively on Cypripedium, Spiranthes, and
Platanthera (among others) for a number of years.
131
5th ANNUAL NORTH AMERICAN NATIVE

ORCHID CONFERENCE
Olympic National Park
Port Angeles, Washington
July 16-20, 2000
Speakers have yet to be arranged, but you can be assured of a varied and
interesting program.
Field Trip highlights will include:
Piperia 4 different species, Epipactis gigantea, Cephalanthera austiniae, Listera caurina,
Corallorhiza mertensiana, Platanthera hyperborea & Platanthera dilatata complexes
And a special trip on the 20th to Lake Elizabeth east of Seattle for Platanthera
chorisiana
And many, many spectacular wildflowers!
Registration fee of $55 per person includes all field trips and conference
sessions
Reservations/registrations should be sent to:
North American Native Orchid Alliance
PO Box 759, Acton, Maine 04001
132
Christenson: ORCHIDS OF MEXICO
ORCHIDS OF MEXICO
Eric A. Christenson
The history of Mexican orchids can be divided

into three distinct periods. The first period was a flurry
of activity during the first half of the 19th century based
on expeditions associated with European centers of
learning. Following this activity there was a lull of more
than 50 years during which little organized collecting or
study took place. The beginning of the 20th century saw
renewed study of Mexican orchids by Americans,
culminating in a full floristic account of the flora in
1951 (Williams, 1951). The third and final phase of
study is the modern period in which Mexican students
have melded extensive field work with taxonomic study
and renewed critical examination of early type
specimens.
The early period:
The first accounts of Mexican orchids comprised

reports on the nature and production of Vanilla. Vanilla
was well established in both horticultural and botanical
literature by the start of modern plant taxonomy in the
mid-18th century.
142
Outside of economic interest in Vanilla, early orchid

collecting in Mexico and the publication of taxonomic
accounts can be divided among five separate groups.
Four of these were standard botanical efforts while the
fifth, and most opulent, was fueled by the first stirrings
of the orchidmania that was to sweep Europe.
The earliest organized collecting of Mexican plants

was by a well funded Royal Expedition from Spain led
by botanists M. de Sessé y Lacasta (1751-1802) and J.
M. Mociño (1757-1820). Sessé actively collected
Mexican plants during the period 1795 to 1804 and they
published the results between 1787 and 1797. Their
contributions to Mexican botany are substantial and are
still being critically investigated (McVaugh, 1998). Their
work had relatively little direct impact on orchid
taxonomy.
The next pair to investigate Mexican plants were P. de

la Llave (1773-1833) and J. Lexarza (1785-1824). They
published two accounts, one in 1824 on general plants
and one in 1825 with an extensive addendum on
orchids. In addition to describing many showy Mexican
orchids such as Barkeria scandens, Euchile citrina,
Rhynchostele cervantesii, and Stenorrhynchos aurantiacus, the
timing of their publication was critical since it came just
before Lindley’s seminal The Genera and Species of
Orchidaceous Plants (1830-1840), generally considered the
starting point of modern orchid taxonomy.
143
The third significant contribution on Mexican orchids
was made by botanist H. Galeotti (1814-1858), who
collected numerous herbarium specimens in Mexico
from 1835 to 1840. Galeotti teamed with French
botanist A. Richard (1794-1852) to produce a treatment
in 1844 based on Galeotti’s collections. This was the
first treatment to include a broad range of genera and
species including many smaller-flowered taxa. Galeotti’s
primary set of collections is conserved in Paris (P) but
many of the corresponding drawings by Richard are to
be found in the Reichenbach Herbarium in Vienna (W);
borrowed by Reichenbach but never returned. Galeotti’s
collections are again being critically studied and the
findings combined with our modern understanding of
species (Salazar, 1994).
About the same time, C. J. W. Schiede (1798-1836)

collected plants in Mexico from 1825 to 1831. Schiede
published very few of his findings but his collections
were described by D. F. L. Schlechtendal and A. de
Chamisso in a series of articles dating from 1830 to
1844 in the important botany journal Linnaea. Schiede is
commemorated in the widely cultivated species
Pleurothallis schiedei and he described Vanilla pompona.
The final contribution during this period was made by

wealthy horticulture patron J. Bateman (1811-1897) who
published the famous folio volume The Orchids of Mexico
and Guatemala in 1843. This magnificent work includes
most of the showy-flowered species and genera in the
region. Not just a pretty picture book, several entities
144
illustrated by Bateman remain enigmatic at this time;
especially the original solid brown-flowered phase of
Cycnoches egertonianum.
The middle period:
For the remainder of the 19th century orchidology

seemed to all but bypass Mexico. The start of the 20th
century, in contrast, saw a dramatic resurgence in
activity through the singular efforts of Swedish expatriot
E. Oestlund (1875-1838). Oestlund, with a group of
associated collectors, began to systematically collect
orchid herbarium specimens. These were funneled to
Oakes Ames (1874-1950) and his colleagues at the
Orchid Herbarium of Oakes Ames at Harvard
University. During the first half of this century, Ames
and his colleagues described numerous new species
from Mexico including a few previously overlooked
showy species such as Euchile mariae.
The culmination of this period was the production of

an annotated checklist by L. O. Williams (Williams,
1951). The efforts of Ames, Williams, and their
colleagues have been unjustly criticized by modern
workers, particularly for their often extremely broad
species concepts. With the available specimens,
equipment, and travel restrictions of the time, they
actually did an admirable job stabilizing names and
providing a useable framework of the orchid flora. It is
easy to forget in the age of paved roads, jet aircraft, and
modern photographic equipment, what a different set of
145
circumstances and opportunities present themselves
today.
The end of this middle period of Mexican orchidology

was the collaboration of G. Pollard in Mexico and R. L.
Dressler in the United States. Their revision of Encyclia,
published in 1974, transformed our understanding of
one of the most taxonomically difficult and species-rich
genera in Mexico. Their effort, strongly influenced for
the first time by extensive field work and knowledge of
living plants in situ, signaled the start of the modern
period. Both taxonomic methodology and philosophy
were changing.
The modern period:
As Dressler and Pollard were completing their work

on Encyclia, a dynamic new group was forming around
the herbarium of the Asociación Mexicana de
Orquideología (AMO). AMO has featured the talents
and contributions of Ed Greenwood, Eric Hágsater,
Frederico Halbinger, Rolando Jimenez, Alicia Rojas,
Gerardo Salazar, Luis Sanchez, Rodolfo Solano, Miguel
Soto, and Roberto G. Tamayo. The AMO group
rejuvenated the journal Orquidea (Mex.). First published
as a single slim pamphlet in the 1940’s, the resurrected
journal began regular publishing in 1971. First given to
species treatments and general orchid activities in
Mexico, the journal quickly pared down to in-depth
taxonomic treatments by AMO Associates and
orchidologists working in Mesoamerica.
146
Recently the format of the journal has changed from

collections of miscellaneous articles to single-theme
books. To date two books have been published on
Lepanthes (Salazar & Soto, 1996) and Laelia (Halbinger &
Soto, 1997) with future volumes planned for Barkeria
(Halbinger & Soto) and Oncidium (R. Jimenez). By
focusing on a single genus augmented with copious
color photographs, these books are more accessible to
growers and other non-taxonomists.
In addition to Orquidea (Mex.), AMO oversees the

production of a loose-leaf illustrated series called Icones
Orchidacearum. Three fascicles of 100 plates each have
been published with one fascicle about miscellaneous
Mexican species and the other two focusing on new
species of Epidendrum.
Floristic relationships
Mexico’s orchid diversity is a combination of endemic

taxa, widespread taxa in common with adjacent
countries, and limited range extensions of orchid genera
more diverse outside of Mexico.
There are only a few endemic genera and, with

exceptions such as Dignathe, Euchile and Hagsatera, these
are all terrestrial species and mostly members of the
Spiranthinae. There are, however, several genera that
although not technically endemic to Mexico find their
147
center of diversity and greatest number of species
within Mexico. In particular Barkeria, Bletia, Laelia (s.s.),
Rhynchostele, and Schiedeella are characteristic of the flora.
Although Mexico shares many taxa with neighboring

regions, recent collecting in Chiapas has recorded
several orchids, such as Stellilabium standleyi, which are
more characteristic of southern Mesoamerica than the
remainder of the Mexican flora.
References:
Bateman, J. 1843. The Orchidaceae of Mexico and Guatemala.

J. Ridgway & Sons, London.
Dressler, R. L. and G. Pollard 1974. El Género Encyclia en
Mexico. Asoc. Mex. Orquid., México, D. F.
Halbinger, F. and M. Soto, 1997. Laelias of Mexico.
Orquidea (Mex.) 15:1-160.
Llave, P. de la and J. Lexarza 1825. Novorum
vegetabilium descriptiones: Orchidianum
opusculum. Fasc. 2:1-43.
McVaugh, R. 1985. Orchidaceae. Fl. Novo-Galiciana 16:1-
363.
---------. 1998. Botanical results of the Sessé & Mociño
expedition (1787-1803), VI. Reports and records
from western Mexico, 1790-1792. IBUG Bol. Inst.
Bot. 6(1):1-178.
Richard, A. and H. Galeotti 1844. Monographie des
Orchidées mexicaines. Comptes Rendu Acad. Sci.,
Paris, 18:497-513.
148
Salazar, G. A. 1994. Identity and synonymy of Notylia
orbicularis (Orchidaceae:Oncidiinae). Lindleyana
9(3):175-182.
Salazar C., G. A. and M. A. Soto A. 1996. El genero
Lepanthes en Mexico. Orquidea (Mex.) 14:1-231.
Sessé y Lacasta, M. de and J. M. Mociño 1787-1791.
Plantae Novae Hispaniae. Naturaleza, ser. 2, 1:1-
184.
---------. 1791-1797. Flora Mexicana. Naturaleza, ser. 2,
2:1-263.
Soto Arenas, M. A. 1988. Listado actualizado de las
Orquideas de Mexico. Orquidea (Mex.) 11:233-
272.
Williams, L. O. 1951. The Orchidaceae of Mexico. Ceiba
2:1-321.
Eric A. Christenson, 1646 Oak Street, Sarasota, Florida,

34236
Eric is a Research Botanist and writes extensively for
many publications including the American Orchids
Society's monthly magazine Orchids.
149
Cameron: PHYLOGENY AND BIOGEOGRAPHY OF POGONIINAE
Cleistes
Isotria
Pogonia
151
PHYLOGENY AND BIOGEOGRAPHY

OF POGONIINAE (VANILLOIDEAE:
ORCHIDACEAE)
Kenneth M. Cameron
The genus Pogonia and its allies, subtribe

Pogoniinae (from the Greek pogon meaning beard), have
been admired by North American naturalists and
studied by professional botanists since Pogonia
ophioglossoides was first described by Linnaeus in 1753.
These orchids are terrestrial and characterized by showy
flowers with abscission layer between perianth and
ovary, bearded or keeled labellum, prominently winged
clinandrium, pollen shed as monads or tetrads, and
fusiform seeds. Vegetatively they may arise from
underground tuberoids or long root-shoots. Their
leaves may be linear, ovate or spatulate, arranged in a
pseudo-whorl or absent altogether. In some species the
foliar venation is reticulate with free vein endings.
Classification and Distribution

The circumscription of Pogoniinae has been
debated by various authors, but most have agreed that
the subtribe includes Pogonia (three species), Cleistes (ca.
60 species), and Isotria (two species) at its core. In
152
addition Duckeella, Pogoniopsis, Triphora, Psilochilus,

Arethusa, Nervilia, and Monophyllorchis have been classified
close to Pogonia at one time or another (e.g., Dressler
and Dodson, 1960; Rasmussen, 1982; Szlachetko, 1995
). Equally debated is the phylogenetic position of
Pogoniinae within Orchidaceae. In many systems of
classification they are positioned close to Vanilla and its
relatives (i.e., subtribe Vanillinae) because of similar
floral morphology and presumed primitiveness
(Dressler, 1981). However, there are no clear characters
uniquely shared between Pogoniinae and Vanillinae
known at this time that support this alliance
(Rasmussen, 1985). Dressler’s (1993) most recent
classification for the orchids placed Vanillinae among
the clades of primitive Epidendroideae while orphaning
Pogoniinae to the category of “misfits and leftovers”.
The most thorough treatment of Pogoniinae was

published by Ames (1922) although he restricted his
discussion primarily to the species of the northeastern
United States. In fact, the geographic distribution of
these orchids extends beyond North America and is
quite a fascinating topic in its own right. The genus
Pogonia is represented by one species, P. ophioglossoides, in
eastern North America that grows in open bogs and
savannas from Florida to Newfoundland, west to Texas
and Minnesota. The remaining two species are found in
Asia, with P. minor being endemic to bogs in Japan, and
P. japonica being found in Japan, China, and Korea. The
wide geographic distance separating these species
demonstrates well the classic eastern North America-
eastern Asia phytogeographic disjunction known for
153
many of our native wildflowers and trees (e.g., Trillium,

Polygonatum, Liriodendron, Panax, Styrax, etc.) as discussed
by Bufford and Spongberg (1983) and others.
An equally interesting geographic disjunction is

demonstrated by Cleistes. With the exceptions of C.
divaricata and the recently segregated C. bifaria, both
from the southeastern United States, and C. costaricensis
which is restricted to Costa Rica, all remaining species
are native to tropical South America. Brazil represents a
center of diversity for the genus with perhaps as many
as 40 indigenous species, and C. rosea is undoubtedly the
most widespread species having been collected
throughout Brazil, French Guiana, Suriname, Guyana,
Colombia, Venezuela, Ecuador, Panama, and Costa
Rica. There are no specimens recorded from Central
America north of Costa Rica or from the Caribbean.
Therefore, Cleistes also displays a rather wide geographic
disjunction; in this case, between North and South
America.
Molecular Systematics
The purpose of this paper is to summarize
recently published information relating to the
phylogenetic and biogeographic patterns among
Pogoniinae. The use of modern DNA sequencing
technology, lightning fast computer hardware, and
sophisticated software for phylogenetic study has
fostered a renaissance in plant systematics. Botanists
now have access to a seemingly limitless supply of
homologous characters (nucleotides) that are essentially
154
free from the selective pressures of the environment.

Direct comparison of the genetic code among different
species has largely confirmed traditional hypotheses of
phylogenetic relationship while challenging others.
These new techniques hold particular promise in
understanding the extremely large and diverse
Orchidaceae. Although orchidologists have been slow
to employ these advances compared to specialists of
other large families (e.g., Poaceae, Asteraceae,
Fabaceae), the value of molecular systematics to
orchidology is quickly becoming evident. Of course,
studies of orchid morphology, anatomy, and cytology
are still as valuable as they always have been, for without
them our gene trees have no story to tell. However, in a
family as diverse as Orchidaceae, it is certain that natural
selection by pollinators and the environment has led to
extremes of convergent and parallel evolution (Dodson,
1962) that may mislead our interpretations of
morphology and ultimately result in erroneous
hypotheses of classification.
The most recently published molecular

phylogeny for the entire Orchidaceae based on
nucleotide sequences of the plastid gene rbcL (Cameron
et al., 1999) showed that the orchids can be subdivided
into at least five subfamilial clades: Apostasioideae,
Cypripedioideae, Orchidoideae, Epidendroideae, and
Vanilloideae. This last subfamily is sister to all
remaining monandrous orchids (i.e., those with a single
fertile anther) and includes Pogoniinae. This same
result was uncovered with a smaller analysis of the 18S
nuclear ribosomal gene (Cameron and Chase, in
155
reviewa). Moreover, these data indicated that

Pogoniinae includes Duckeella, Cleistes, Isotria, and
Pogonia. It does not include Monophyllorchis, Triphora,
Arethusa or Nervilia, all of which are members of the
“lower Epidendroideae”. DNA from the rare, myco-
heterotrophic genus Pogoniopsis has not yet been
obtained; it may ultimately be found to belong in
Pogoniinae as well.
Morphological Studies
Expanding upon the results of the studies
mentioned above, Cameron and Dickison (1998)
studied the foliar architecture of select species from
Vanilloideae including eight species of Pogoniinae.
They found that leaf morphology and anatomy were
quite variable among the genera and that a clear
evolutionary continuum was evident among these extant
orchids. Leaves from Duckeella and Cleistes represent the
ancestral condition of parallel primary veins with evenly
spaced secondaries. In Isotria, the primary veins are also
parallel, but the distribution and orientation of the
secondary veins results in a weakly reticulate pattern.
Finally, truly reticulate leaf venation with distinct free
vein endings are found in Pogonia representing the most
derived condition in the subtribe. These patterns shed
further light on how reticulate leaf venation may have
evolved in other monocotyledons.
156
Seed morphology within Vanilloideae has also

been examined recently (Cameron and Chase, 1998).
Seeds from these taxa are among the most varied and
unusual in all of Orchidaceae. They may be spherical
and sclerified as in Vanilla, elongate and filamentous as
in Lecanorchis, or variously winged as in Eriaxis,
Epistephium, and Galeola. The seeds of Pogoniinaeare
more typical of the family, however, being fusiform and
tunicate. Differences among genera and species are
evident, nevertheless, and of potential systematic value.
For example, the seeds of Cleistes divaricata were found
to be most similar to those of Isotria and Pogonia. They
are markedly different in size and shape from tropical
species of Cleistes and may indicate a stronger
relationship of that taxon to its temperate cousins than
to its tropical congeners.
Phylogeny and Biogeography

Finally, in order to address interspecific
relationships within Pogoniinae, Cameron and Chase (in
reviewb) sequenced the internal transcribes spacers (ITS)
of the 18S-26S nuclear rDNA repeat. Combining these
data with the rbcL sequences and 48 morphological
characters produced cladograms of highly-supported
interspecific relationships for Pogoniinae. In the most
derived position of the tree sits Pogonia. Specifically, the
two Asian species are most closely related and P.
ophioglossoides is sister to them. Sister to Pogonia in most
trees are Isotria and Cleistes divaricata which are closely
related to each other. Following along, one finds that
the two tropical species of Cleistes sampled are closely
157
related and, as a pair, are sister to the temperate clade.

Finally, the enigmatic genus Duckeella with its nearly
actinomorphic, yellow perianth is sister to all remaining
Pogoniinae. Thus, according to these combined data,
the genus Cleistes is not monophyletic. It may be that all
the North American and Asian taxa should be lumped
back into Pogonia. Alternatively, Cleistes divaricata and C.
bifaria must be given a new genus name. Cameron and
Chase (in reviewb) suggested that neither action be taken
at this point in time until further sampling within Cleistes
is completed. A summary of these results is depicted in
Figure 1.
With a robust phylogeny in hand, the issue of

historical biogeography in Pogoniinae could be
addressed. Cameron and Chase (in reviewb) suggested
that the group originated in South America during the
late Cretaceous (ca. 80 million years ago). Here it
underwent speciation and diversification, and ultimately
migrated to the southeastern corner of North America
via a long distance dispersal event sometime in the early
Tertiary (ca. 50 mya) when those two continents were
much further apart than they are today and before the
existence of Central America. The proposed timing of
these events was based on calculations of percent
genetic divergence from the rbcL gene between extant
species pairs coupled with published estimates of
divergence time for herbaceous flowering plants (i.e., a
“molecular clock” approach). With the ancestor of
North American Pogoniinae firmly established on this
continent, northward migration and speciation took
place in a rapidly changing Oligocene-Miocene
158
environment (we still see the “footprints” of these

ancient migrations in the current distributions of Cleistes
divaricata, Isotria, and Pogonia ophioglossoides as depicted in
Figure 2). During the Miocene the genus Pogonia would
have had a widespread range across most of northern
North America and northeastern Asia. The climate in
this region was much more mild than it is today.
During the late Tertiary and early Quaternary increased
drying and cooling of central North America, ultimately
resulting in glaciation through the middle of Pogonia’s
range, would have fragmented the large population into
two, disjunct smaller ones that survived in eastern
North American and eastern Asian refugia. The
speciation of P. japonica and P. minor in Asia was the final
result of this geographic isolation.
Conclusions
Molecular studies of vanilloid orchids, including
Pogoniinae, have provided compelling evidence to help
answer many perplexing questions concerning these
beautiful and fascinating plants. Many assume that
North American orchids are fully studied and that there
is nothing more to learn from them. The studies
discussed in this paper confirm that this is far from the
truth and that many of our native orchids have a great
deal to teach us about plant biology and evolution.
Unfortunately, these are among the most threatened
plants in North America and efforts to conserve
particularly vulnerable species (such as Isotria medeoloides -
- a member of Pogoniinae and one of our most
endangered orchids) must be put into place
159
immediately. Further research is underway to learn

more about these bearded orchids, especially the
tropical members, before deforestation and wetland
destruction take them away from us forever.
Literature Cited
Ames, O. 1922. A discussion of Pogonia and its allies in the
northeastern United States with reference to extra-limital
genera and species. Orchidaceae 7: 3-38.
Bufford, R. J. and S. A. Spongberg. 1983. Eastern Asian-eastern
North American phytogeographical relationships –
history from the time of Linnaeus to the twentieth
century. Ann. Missouri Bot. Gard. 70: 423-439.
Cameron, K. M. and M. W. Chase. 1998. Seed morphology of
vanilloid orchids (Vanilloideae: Orchidaceae). Lindleyana
13: 148-169.
Cameron, K. M. and M. W. Chase. in reviewa. 18S rDNA
sequences confirm the subfamilial status and
circumscription of Vanilloideae (Orchidaceae). In K.
Wilson (ed.), Proceedings of Monocots II, CSIRO Press,
Sydney, Australia.
Cameron, K. M. and M. W. Chase. in reviewb. Phylogenetic
relationships of Pogoniinae (Vanilloideae, Orchidaceae):
an herbaceous example of the eastern North America-
eastern Asia phytogeographic disjunction. Journal of Plant
Research.
Cameron, K. M. and W. C. Dickison. 1998. Foliar architecture of
vanilloid orchids: insights into the evolution of reticulate
leaf venation in monocotyledons. Bot. J. Linn. Soc. 128: 45-
70.
Cameron, K. M., M. W. Chase, W. M. Whitten, P. J. Kores, D. C.
Jarrell, V. A. Albert, T. Yukawa, H. G. Hills, and D. H.
Goldman. 1999. A phylogenetic analysis of the
160
Orchidaceae: evidence from rbcL nucleotide sequences.

Amer. J. Bot. 86: 208-224.
Dodson, C. 1962. The importance of pollination in the evolution
of the orchids of tropical America. Amer. Orchid Soc. Bull.
31: 525-554.
Dressler, R. L. 1981. The Orchids, Natural History and Classification.
Harvard University Press, Cambridge, Massachusetts.
Dressler, R. L. 1993. Classification and Phylogeny of the Orchidaceae.
Cambridge University Press, Cambridge, Massachusetts.
Dressler, R. L. and C. Dodson. 1960. Classification and
phylogeny of the Orchidaceae. Ann. Missouri Bot. Gard. 47:
25-68.
Rasmussen, F. N. 1985. Orchids. In R. M. Dahlgren, H. T.
Clifford and P. F. Yeo (eds.) The Families of the
Monocotyledons, Springer Verlag, Berlin.
Rasmussen, F. N. 1982. The gynostemium of the neottioid
orchids. Opera Bot. 65: 1-96.
Szlachetko, D. L. 1995. Systema Orchidalium. Fragmenta Floristica
et Geobotanica Supplementum 3: 1-152.
Kenneth M. Cameron
The Lewis B. and Dorothy Cullman Program for Molecular Systematics
Studies, The New York Botanical Garden, Bronx, NY 10458-5126 Email:
kcameron@nybg.org
Acknowledgments
Gratitude is extended to Mark Chase and William Dickison
for guidance during the preparation of this study and to Gustavo
Romero, Eric Christenson, and Katsuhiko Kondo for their role
in obtaining plant material. Financial assistance was partially
provided by the Lewis B. and Dorothy Cullman Foundation.
161
Figure 1. Phylogenetic relationships within Pogoniinae and

the placement of the subtribe and suspected relatives
among clades of Orchidaceae. The tree represents a
generalized summary of results based on published studies
of molecular and morphological analyses.
162
Figure 2. The approximate geographic distribution of

Pogoniinae genera in North America. Pogonia is also
found in eastern Asia, and the great majority of Cleistes
species are native to South America.
163
Sampliner: KEYING OUT A RED MITSUBISHI
KEYING OUT A RED MITSUBISHI

or the Trials, Travails, and Tears of an Orchid Hunter
Tom Sampliner
Ah, yes. Escaping the Cleveland cold for four

days of field trips and two of lecture; in Florida no less
and upon one of my most favorite topics: native
orchids. Despite the initial shock of waiting in line at
Alamo for over an hour just to get my rental car, events
really went rather smoothly for the first two days of
field trips in northern Florida. Granted conditions were
unusually dry to the point we were not seeing all we
hoped for. The fact the state was under alert and
National Guard troops were being called out to assist in
fire fighting efforts. Still we were seeing new to many of
us species and enjoying the warmth and scenery of the
state.
164
Saturday arrived before we knew it and the first of

two days of lecture presentations at the University of
South Florida. Featured were well known experts both
within Florida and from the national scene.
At lunch break Saturday, I decided to employ my

usual procedure of obtaining take out from the nearest
grocery and picnicking at one of the several metal
grillwork style tables and benches sprinkled through the
campus. I recall laying all my conference notes, packet,
maps and other material upon the table. Cute,
precocious gray squirrels were everywhere. So tame
were they, not to mention hopeful for a handout, that
they literally shared both bench and table. We
entertained each other thoroughly. They did not
however, seem interested in my neptune salad; so much
for their taste. Anyway, upon finishing, I folded up the
trash and took it to the nearest container to make my
deposit. I had time to return to the car, a red Mitsubishi
eclipse, and shift some belongings, maybe even take
some photographs on the attractive campus.
Upon reaching the car I reached into my pocket

to retrieve the key; no key. I searched and researched
each pocket. I look into the car but see no key visible.
Next I return to the picnic table and search the tabletop,
the ground beneath and surrounding paths I had to
have taken. No key. I go over to the waist high garbage
container, reach in and retrieve the remnants from
lunch; no key. I remove the garbage can lid and remove
by hand all contents from the can but still find no key.
165
This is not looking good. I return to the lecture

hall and search all around the registration desk and the
seat I occupied during the morning as well as rows up
and downward in the theatre style lecture hall. I am
beginning to get a sinking feeling in the pit of my
stomach and it was not from lunch.
The only thing to do was to call Alamo; I was

about to discover they were well named, for it was fast
becoming my last stand. When you call a big company
these days you first go through the menu and only then
begin the mandatory stay on hold for an eternity until
something semi-human comes on line. That
accomplished, I explained my plight to the gentleman.
He said if they had to come out it would take quite a
while and I would be charged for their having to
duplicate a key. He suggested that if I was an AAA
member, I should call for emergency road assistance
and that they would probably be here even faster than
Alamo could and that I would not be charged anything.
It sounded reasonable so I hung up and called AAA. As
you already know, menu, stay on hold, etc. until you get
a human It being very hot early in their spring, 90's, they
were very busy but would be out in around 45 minutes.
Be by the car.
As promised, the independent contractor, who is

also a locksmith, arrived and opened the car. We
searched all over but found no key insider was informed
that car keys, especially rentals, cannot just be
duplicated by anyone. You need a key's code from
which the cuts in the metal can be guided. Is this
166
anything like a dichotomous key? Probably not.

Anyway, I was told I must recall Alamo and get the
code then I could have AAA come back to make a new
key. It sounded very logical. Of course, even though I
timed the making of my calls to coincide with an
afternoon break, I was beginning to miss some lectures
I wanted to hear.
I called Alamo, menu, mandatory hold time,

then human. I explained the scenario; no - customers
can not have a key code. Thinking quickly and
becoming frustrated, yeah it was a dichotomous key
after all, I asked if they would give it to AAA if they
were to call; Alamo said yes.
I recalled AAA and once through, the

representative conferred with a supervisor and they
agreed to try to get the code for me. Their wait time was
now estimated to be over one hour- by now that meant
bye-bye dinner for me.
The same jovial guy drove back up about l 1/4

hours later. Home base had not obtained any code nor
informed him. However, he rapidly employed his cell
phone and home base called Alamo and began to argue
on my behalf. We knew it was a dispute because after
some 15 minutes of back and forth we were told the
code would be given if we supplied the car's VIN
number from inside the vehicle. That done we were
given the code, wrong the first time naturally, and then
correct. It took a couple manufacturing attempts before
a workable key was made. Meanwhile, some friends
167
attending the conference brought me back a sandwich

for dinner.
AAA could not have been more understanding

throughout. The driver explained that in Florida, expect
this kind of treatment from all the rental agencies; it's a
seller's market. Before we were sure we were getting the
code, we joked about telling Alamo to just come take
their rental back.
Thinking back to my hour plus wait in line just to

get the car, I was a little envious of those who, upon
seeing the line as they got off the courtesy van and
learned of the wait time, simply went back to the airport
to rent from one of the non-discount companies at a
higher price.
So this, dear readers is how one keys out a red

Mitsubishi. Next time think I'll come equipped with
peanuts - they key out so much easier.
Tom Sampliner, 2651 Kerwick Rd., University Heights, Ohio 44118

Tom is both an excellent botanist and natural history
photographer and lead trips to various parts of North America.
He had regularly contributed his adventures to the Journal.
168
Brown: A STRIKING NEW COLOR FORM OF SACOILA
LANCEOLATA
RECENT TAXONOMIC AND

DISTRIBUTIONAL NOTES FROM
FLORIDA 2.
A Striking New Color Form of Sacoila lanceolata
Paul Martin Brown
Sacoila lanceolata var. lanceolata (Aublet) Garay, the

red ladies-tresses or leafless beaked orchid, is a not
infrequent member of the flora of central Florida. It
can been seen along the major highway right-of-ways
from the Orlando-Tampa area southward to Fort
Myers. Beyond this area it occurs sparingly as far north
as Bradford County, with a historical record for Baker
County, and south to Dade County. A large
concentration of this species occurs along Florida's
Turnpike near the Yeehaw Junction exit and the Fort
Drum Service Area. While monitoring the populations
in this area we observed both the typical red (which
actually varies from coral to pink to brick red) and a few
of forma albidaviridis. Nearly 5000 flowering stems of
the red plants were counted in May of 1999 in this area.
Scattered in 8 sites along the turnpike were a total of 67
flowering plants with bronzy-gold flowers and the lips a
169
LANCEOLATA
pale yellow with a rose overlay. This new and striking

color form is described below.
Sacoila lanceolata (Aublet) Garay var. lanceolata

forma folsomii P. M. Brown forma nov.
Forma floribus aeneus-ochraceus cum erubescens

conspeciebus diversa
Differing from other forms of the species by it coppery-
golden yellow flowers with palest pink overlay.
TYPE: UNITED STATES: Florida, Okeechobee

County, along the Florida Turnpike. (Holotype. Brown
99-527, May 29, 1999. FLAS) Photo: North American
Native Orchid Journal 1999. 5(2): plate 3
PLANTS often grow within populations of typically

'red' colored Sacoila lanceolata var. lanceolata and
occasionally with the green and white forma albidaviridis
Catling & Sheviak. In a brief, but critical note Catling &
Sheviak (1988) reviewed the status of color forms of
Sacoila lanceolata var. lanceolata and at that time published
forma albidaviridis to satisfy a discrepancy with that of
var. luteoalba (Rchb.f.) Luer that Luer had used in his
work (Luer, 1972). The presence of this new unusual
color form raises the questions again as to the correct
status of the white and green flowered plants. Luer saw
them as quite distinct from the red flowered plants and
therefore used the rank of variety. Catling and Sheviak
pointed out that within the range of the species many
local races occur as the species is apomictic and
novelties reproduce as pure lines. The first impression
170
LANCEOLATA
of forma folsomii is that they are hybrids between typical

'red' lanceolata and the white and green albidaviridis. If
albidaviridis is a true 'alba form' and apomictic how
would this be possible? Or are there sexually
reproducing plants within 'red' lanceolata or albidaviridis to
be found in Florida? All of the folsomii are found within
a 10-mile stretch of the turnpike and in one instance the
largest site is exclusively folsomii (this year). It is well
known that not all plants within a given population of
Sacoila flower each year. At this time it is thought that it
would be best to designate folsomii as forma rather than
to hypothesize at another taxonomic level.
This situation must make us re-examine other

populations and carefully look at forma albidaviridis.
Luer (1972) very carefully noted several differences
between what we now know as forma albidaviridis and
typical 'red' lanceolata. Some of the differences do not
bear up when examining many populations. The vast
majority of forma albidaviridis are at least superficially
different morphologically from the typical 'red' plants
with the flowers held at a sharper angle and the plants
being strongly three-ranked. Catling & Sheviak (1993;
Catling, 1987), state that throughout the range of the
species through Central and South America these traits
may also be present in the 'red' forms and the species
presents an even more interesting situation. Forma
albidaviridis had been observed to have uniformly smaller
ovaries and narrower floral parts than the 'red' lanceolata
flowers
171
LANCEOLATA
If individual plants are facultativly apomictic, and

some flowers function sexually, or, if there is a non-
apomictic race of either forma albidaviridis or 'red'
lanceolata then it is entirely possible that hybrids between
the races could occur and that their progeny would also
be apomictic. Subsequently those flowers that produce
seed therefore could result in large numbers of the gold-
colored plants. Future studies of the pollination
mechanisms of these plants may aide in determining the
correct status of forma albidaviridis and forma folsomii
(Sheviak, pers. comm).
Small populations of Sacoila lanceolata occur

scattered throughout much of Florida but in these
smaller populations (1-50 plants) no forma albidaviridis
or forma folsomii were noted. Only in the larger
population areas (300-1000+ plants) were the color
variations seen. Due to extensive road reconstruction in
the Orlando area, few other large populations of Sacoila
still are extant in Florida and they have been carefully
searched for additional forma folsomii plants. One
flowering plant of forma folsomii was found on 5 June
1999 in northern Collier County. It was growing within
a group of predominately forma albidaviridis and a few
typical 'red' lanceolata.
ETYMOLOGY: name for Stanley Noyes Folsom

(1933 - ), artist and botanical illustrator who first
noticed the plants during a survey of the area.
Acknowledgments:
172
LANCEOLATA
I thank Charles J. Sheviak (NYS) for his expertise and advise in

the preparation of this paper.
Literature Cited:
Catling, P.M., 1987. Notes on the breeding systems of Sacoila
lanceolata (Aublet) Garay (Orchidaceae). Ann. Missouri Bot.
Garden 74:58-68.
Catling P.M. & C.S. Sheviak. 1993. Taxonomic notes on some
North American Orchids. Lindleyana 8(2): 77-81.
Luer, C.A. 1972. Native Orchids of Florida, New York Botanical
Garden, Bronx.
Folsom's golden ladies'-tresses

Sacoila lanceolata var.lanceolata
forma folsomii
173
Folsom: HOW IT ALL CAME ABOUT
HOW IT ALL CAME ABOUT
Stanley N. Folsom
Paul Martin Brown and I had gone down to West

Palm Beach where Paul was to give a talk about his
native orchid project here in Florida. On the way back
Paul saw a large stand of the red ladies'-tresses, Sacoila
lanceolata var. lanceolata, along the Florida turnpike north
of the Fort Drum Service Area. We pulled over and did
a short exploration and realized there were many plants
only in bud. We agreed that someone should be notified
about them. Before we left the spot, I looked over the
toll ticket to find out if there was any information
printed on it to help us contact the proper authority. I
found a toll free number for the turnpike and copied it
down. We also discovered that we could get our ticket
stamped at any rest area to make a U-turn on the
turnpike. As we continued our drive toward home we
spotted several other sites and Paul, who was spotting,
noted the mileage markers near the sites. Another large
stand was seen nearly in downtown Orlando.
The next day, Paul was able to reach the proper

people and set up a time to show them where the plants
were near the Fort Drum Service Area. On the
174
scheduled day, we bundled our equipment and our two

Pomeranians into the wagon and set out. I liked to drive
the initial leg of the trip first so that if I get weary, Paul,
who has more energy, can relieve me but because we
saw the plants further into our trip, I let Paul drive the
first leg. As we neared the area where we had started to
see the orchids, I took over the driving. Paul could spot
orchids from the passenger side more easily. He noted
their position from the mileage markers but we did not
stop to check any sites, as we would be late for our
meeting with the turnpike people. Paul spotted some of
the white and green form, forma albidaviridis, of the
Sacoila lanceolata. I asked if there were any yellow forms
of the orchid present in the United States. Paul replied
that there were none ever found. He said that it was a
mistaken report that was never verified. I said that if
there were yellow ones I would have sworn I saw some
back near the Orlando area. I dismissed my
observations as probably being particularly lush plants
of dock, which was abundant along the roadside.
When we reached the rest area and met our

turnpike people, Lauren and Ben, we got our toll ticket
stamped, walked the dogs and set off for the large stand
of Sacoila lanceolata. As soon as we arrived at the area,
Paul was out of the car and showing the people the
plants. Since they stand about 12 to 24 inches or better
in height and there were several hundred of them in
bloom, they were easy to see. I walked the dogs again
and headed back east as they headed further west up the
colony. I found two beautiful specimens growing in
close proximity that were in perfect condition. I walked
175
along further until the plants ceased before turning

back. By this time Paul and his companions had reached
the other end and were spending a seemingly inordinate
amount of time up there. I thought, "They must have
found something different."
As Paul came back toward the car he announced

that he had found a new color form for the Sacoila
lanceolata and it was gold. I went up to inspect the find
and surely enough, it was a tawny bronze, sort of an
antique gold with a pale wash of rosy orange to it. Near
the gold plants there were many of the red form and
one white and green form. There were 17 gold plants in
all. A very exciting find!
The two people from the turnpike enlisted Paul

to do a survey of the Sacoila lanceolata plants along the
turnpike. They gave us a yellow flashing beacon to put
on the roof of the car and told Paul to wear his orange
vest while doing it. They also gave us several other
cautions about doing the survey in safety. As we drove
home late that afternoon, we noted sites as we drove
past them and tried to locate the gold ones I had seen
on the way down. We couldn't see any sign of them. We
also saw that the large stand of Sacoila lanceolata near
Orlando we saw the week before had been mowed
down leaving only a tight green lawn. We decided that
we had better do our survey as soon as possible in case
the rabid mower struck any more areas.
Consequently, we set out early on that Saturday

to comb the highways in hopes of finding more gold
176
form plants and seeing if we could verify my

observations. We could not relocate my original sites
and have concluded that they were indeed probably
dock. We did a very thorough search of the north
bound and south bound sides of the road all the way to
Fort Pierce. Paul noted mileage and numbers and kinds
of plants on a check sheet that I had devised, plus any
notes that were important. I did all the driving in the
critical areas. In all we spent 12 hours in our initial
foray. The Pomeranians were well behaved, the weather
cooperated, and only darkness and mowed stretches
thwarted our efforts. Just past the Fort Drum rest area
going south I spotted the illusive gold form of the
orchids. We had quite a stand of them and two plants
that took the configuration of the red form rather than
the three-ranked configuration of the white and green
form that all the other gold forms displayed. I was
elated because Paul had wanted to name the new form
after me since I had supposedly seen it first. I felt that
my sitings were highly suspect and I didn't deserve to
have the form named after me. After discovering my
own stand, I felt that I could let the plants carry my
name in the description. It was very thrilling to finally
have my name on a description after years of looking
for orchids.
Our efforts that day yielded a total of 8 sites for

the gold form with 67 plants and 16 sites for the white
and green form with 93 plants. There were areas where
there were over a thousand plants of the red form. We
will be continuing to monitor the plants and check out
other sites.
177
Stanley N. Folsom, 10896 SW 90th Terrace, Ocala, FL 34481

Stan's line drawings have been a regular feature of this Journal
since its inception, as well as an occasional article.. He is an artist
and watercolorist of note and has co-authored Wild Orchids of the
Northeastern United States with Paul Martin Brown. Together they
are currently working on a field guide to the orchids of Florida.
178
POLLINATOR OF CLUSTERED LADYS-
SLIPPER ORCHID, CYPRIPEDIUM
FASCICULATUM, (ORCHIDACEAE) IN
OREGON
C. S. Ferguson & K. Donham
Cypripedium fasciculatum, clustered lady's-slipper,

(Orchidaceae) is one of three endemic Cypripedium
species in the western United States (Knecht 1996). In
Oregon, the USDI Bureau of Land Management lists C.
fasciculatum, the Clustered lady's slipper orchid, as sensitive. A
number of researchers assert that pollinators are
necessary for good seed set in C. fasciculatum, and the
unique floral morphology suggests a close evolutionary
relationship with a particular pollinator. Insect
pollinators of other Cypripedium species include bees
(Andrenidae, Apidae, Anthophoridae, Halictidae,
Megachilidae) and flies (Syrphidae); however, the
pollinator(s) of C. fasciculatum have been to date
unknown (Catling & Catling 1991). The purpose of this
research was to determine the pollinator(s) of C.
fasciculatum in Southwest Oregon populations.
179
Field studies were conducted between April and
June of 1998 and 1999. Insects were collected by
various methods from the Cypripedium sites. Cypripedium
floral phenology, pollen condition, and flower/insect
associations were also measured. During 1998, an insect
carrying a pollen smear on the dorsal side of its thorax
was removed from a Cypripedium flower as it was exiting
the posterior labellar opening. The insect was collected
intact and the pollen was confirmed as Cypripedium
pollen. During the current field season we have
collected nine additional insects from field sites in
Jackson and Josephine Co., OR with Cypripedium pollen
on their thorax. In all cases, the insect has been
identified as a wasp (Hymenoptera, Family Diapriidae,
Subfamily Belytinae). Species identification of the wasp
is being done by Dr. Lubomir Masner (Ottawa Canada).
This is the first record of a parasitic Diapriid wasp as a
pollinator.
The Diapriids are a very large family of relatively

obscure wasps with over 300 species in North America
(Borror et al. 1989). Although they are a large family
and distributed worldwide, very little is known about
their life history. Past studies on their biology suggests
that most Diapriids are endoparasitoids that develop in
Dipteran pupae (Gauld & Bolton 1988). The Subfamily
Belytinae have been successfully reared from both
Mycetophilidae and Sciaridae (dark-winged fungus
gnats) (Gauld & Bolton 1988). Both families of fungus
gnats have been collected from the Cypripedium field
180
sites and both have been observed closely associated
with the plants. Besides serving as a host to Diapriid
wasps, Mycetophilids have also been implicated as
pollinators of other orchids (Mesler et al 1980), and may
prove to be important in pollen transfer in C.
fasciculatum. Therefore, both insects may be important
in the successful pollination of C. fasciculatum and
additional life history information on both groups needs
to be elucidated. We hypothesize that floral cues may
attract both the fungus gnats (Mycetophilidae, Sciaridae)
and the parasitic wasp (Diapriidae) and hope to clarify
this relationship in future studies.
References Cited
Borror, D., Triplehorn, C., & Johnson, N. 1989. An Introduction to the Study of
Insects. 6th ed. Harcourt Brace. Orlando, FL.
Catling, P.M. & V.R. Catling. 1991. A synopsis of breeding systems and
pollination in North American Orchids. Lindleyana. 6(4):187-210.
Gauld, I. & B. Bolton [eds]. 1988. The Hymenoptera. Oxford University Press.
Knecht, D. 1996. The reproductive and population ecology of Cypripedium fasciculatum
(Orchidaceae) throughout the Cascade Range. MSc. Thesis. Central
Washington. University, Ellensberg, WA.
Mesler, M.R., Ackerman, J.D. & K.L.Lu. 1980. The effectiveness of fungus
gnats as pollinators. Amer. J. Bot. 67(4):564-567.
C. S. Ferguson, Biology Department, Southern Oregon University, Ashland

OR 97520
K. Donham, 3776 Devils Garden Rd, Medford, OR 97504
This research was supported in part by the Bureau of Land Management,
Medford District, and the US Department of Fish and Wildlife.
181
Latham: A STRATAGY FOR COPING WITH RARITY IN CYPRIPEDIUM
FASCICULATUM
A STRATEGY FOR COPING WITH

RARITY IN THE ORCHID
CYPRIPEDIUM FASCICULATUM
Penelope A. Latham
The clustered lady’s slipper (Cypripedium

fasciculatum Kellogg ex. Watson) is a small terrestrial
orchid that occurs in seasonally dry forest habitats of
the northwestern United States. Because the plants are
rare and existing populations generally occur on forest
lands managed for timber or other resource values, they
may be at risk. The species is currently managed
according to special criteria in the Record of Decision
of the Northwest Forest Plan (1994). In response to
the pressing need for better information with which to
manage this species in southwest Oregon, a long-term
study of the autecology, habitat characteristics,
successional status, and population dynamics of the
clustered lady’s slipper was initiated by the
Cooperative Forest Ecosystem Research program at
Oregon State University in 1998.
Cypripedium fasciculatum is one of three Cypripedium

species endemic to the northwest. The other two
species, mountain lady’s slipper, Cypripedium montanum,
182
FASCICULATUM
and California lady’s slipper, Cypripedium californicum,

are also rare although both of these species occupy
more specialized habitats than the clustered lady’s
slipper. Relatively inconspicuous compared to these
showier species, the inflated slipper petal is a delicate
pale green with streaks of brownish purple on the lateral
edges (Figure 1). The labellum is surrounded by five
similar tepals: two lateral petals and three sepals (two of
which are united beneath the labellum). The tepals vary
in color from a striking deep red to a dull brownish red
or even a yellowish-green. The flowering habit ranges
from a single flower on a single aerial stem to several
flowers (5-6 or more) on each of 4-5 aerial stems
grouped at the anterior end of a short underground
rhizome (generally 3 - 5 cm for larger vegetative to
mature individuals). Individual plants usually occur in
close proximity to each other. The habit of grouping
flowers and stems has given rise to the name clustered
lady’s slipper (Elliman and Dalton 1995).
Rarity, Population Structure, and Pollination
Several different forms of rarity exist which
depend upon the unique combination of frequency and
abundance exhibited by a species (Rabinowitz 1981).
Cypripedium fasciculatum occurs both infrequently and in
low abundance. Despite the fact that Cypripedium orchids
are among the most prized of North American
terrestrial orchids, little is known about their autecology.
While most Cypripedium species are insect-pollinated
(Cribb 1997, Catling and Catling 1991), the pollinator of
C. fasciculatum has remained a mystery until recently [See
Ferguson and Donham (this issue)]. However, at least
as intriguing as the identity of the pollinator is the
183
FASCICULATUM
question of how a species that occurs infrequently and

in low numbers manages to consistently attract an insect
pollinator. It is well-documented that many rare species
are either self-pollinating or need more focused pollen
vectors than more common species (Gaston and Kunin
1997). Compounding the problem is the fact that
Cypripedium orchids are well-known for their use of
deceit to lure insect visitors into their trap-like labellum
and do not provide a reward for services rendered
(Cribb 1997).
One hypothesis being testing is that the habit of

clustering flowers, stems and individuals is adaptive. If
so, then greater fruit production should occur in
populations showing those characteristics. Elliman and
Dalton (1995) state that single stems are atypical and
that the usual subpopulation cluster is three to ten stems
clumped in one spot. Unfortuately, little quantitative
historical data exists regarding the distribution and
abundance of the clustered lady’s slipper. Whittaker’s
(1960) data from the Siskiyou Mountains indicates that
populations may previously have been more infrequent
but were composed of greater numbers of stems.
However, because only the most general frequency and
density description is available (stems/.1-hectare), this
conclusion should be regarded with caution.
Analysis of over 430 sighting reports in

southwest Oregon has shown that 77% of C. fasciculatum
sightings on federal lands consist of populations of 1-10
stems and is consistent with Elliman and Dalton’s
(1995) observation. Observation of over 60 populations
184
FASCICULATUM
in southwest Oregon, clearly shows that a range of

population structures exist even though the majority of
current populations are small. Some of these sites have
revealed greater numbers of stems upon visiting them
earlier in the season. Whether these stems are residual
or colonizing is unclear. Size classes are currently being
developed and tested that will assist in clarifying
population dynamics. The question remains, however,
whether floral clustering is an adaptive strategy for the
clustered lady’s slipper.
Another hypothesis is related to Cypripedium

fasciculatum phenology. In view of the preponderance of
small populations, a mechanism for ensuring pollination
of C. fasciculatum flowers might be to flower
concurrently with other spring species. Hypotheses
found in the literature cover a range of situations.
Divergent flowering patterns of sympatric species have
been explained as a means for different species to
compete for limited pollinator services. Convergent
flowering patterns have been explained as adaptations
for tracking seasonal changes in pollinator abundance or
for satiating seed predators. Intraspecific synchronous
flowering has been suggested as a strategy to attract
pollinators during brief flowering periods, whereas,
steady-state flowering may be advantageous for
spreading the risk of uncertain pollination (Rathcke and
Lacey 1985).
Methods, Observations, and Results

To explore these hypotheses, I used data
gathered from 163 mapped and tagged Cypripedium
185
FASCICULATUM
fasciculatum stems (77 of which were flowering) at four

sites in the 1998 pilot study. The number of stems with
fruits were tallied by flower number per stem across all
sites. Fruit production was also compared by flower
number per stem. Flower and fruit production, and the
number of stems with multiple flowers were compared
at each site.
In 1999, a phenology study was initiated at 15

sites along an elevation gradient in five different regions
of the Siskiyou and southern Cascade Mountains.
Weekly observations are being made on over 100
flowering stems. By the end of the season, we will have
data on approximately 500 mapped individuals at 20 -
30 sites. Observations included in this preliminary
report are primarily from three sites in one
environmental region. Total population sizes at these
sites vary from 17 to 83.
Data from the pilot study indicated that fruit production
for stems with single flowers was only 14.7% compared
to 55.6% for stems with more than one flower. Fruit
production for the 4 flower per stem group did not
continue to increase as did the previous two groups
(Table 1); however, the number of stems in our data
with 4 flowers per stem was small compared to the
other groups. When fruiting is compared across sites,
the results are less clear (Table 2). Site 3, a large
population with the second highest fruit production had
the lowest percentage of stems with more than one
flower. Site 4, a small population with the lowest fruit
production had the highest percentage of stems with
more than one flower. It may be that greater numbers
186
FASCICULATUM
of flowers per stem are more important for populations

with fewer numbers of stems. Certainly, the
relationships are complex and clues to the floral strategy
of Cypripedium fasciculatum will not come easily.
Hopefully, a more complete data set will be revealing.
Preliminary observations from this year’s

phenology study have produced very interesting results.
First, confirming last year’s observations, C. fasciculatum
flower buds are present in the leaves as they emerge.
Apparently, the buds are not fully developed at this
time, because their display takes several days to weeks.
This year, however, as I knelt to closely examine this
small statured plant growing on steep slopes, I
occasionally observed what appeared to be a very tiny
undeveloped bud either in the juncture of the two
subopposite leaves of a “non-flowering” stem or deep
within a new bract on a peduncle with a previously
existing flower (Figure 2). Thinking at first, that these
structures were browsed or aborted buds, I followed
them closely and observed them growing in size. The
larger buds (2-3 mm) eventually take on the dark
reddish coloration of mature flower tepals. Closer
observation of mature C. fasciculatum inflorescences
revealed that flowers are quite variable in size. Could it
possibly be that C. fasciculatum flowers take several years
to develop? Although this does not seem to be a very
productive strategy, my observations of this species
indicate that growth and development occurs rather
slowly. For example, one non-flowering individual with
an undeveloped bud at the leaf juncture was estimated
to be approximately 22 years old based on the number
187
Latham: A STRATAGY FOR COPING WITH RARITY IN
of roots that were present when excavated [Latham,

unpublished data; Also see Curtis (1943)]. These
diminuitive buds will be followed closely for the next
several years to determine their ultimate fate.
With respect to the convergent flowering

hypothesis, preliminary observations suggest that the
clustered lady’s slipper may have evolved a specific
phenological niche so that it competes for pollinators
with few (if any) other species. Thus, the opportunity
for pollinators to be carrying pollen from heterospecific
flowers before visiting a C. fasciculatum flower is reduced.
More frequent development of fruits on stems with
greater numbers of flowers would seem to reinforce this
strategy. Thus, a successful pollination strategy for the
clustered lady’s slipper might be to aggregate flowers in
space and time, and to select that time such that few
other flowers are available to distract potential
pollinators. Such a strategy would depend upon the
pollinator being both abundant and regionally
distributed.
At present, the clustered lady’s slipper remains an

enigma and a challenge to resource managers and
conservationists, but one whose secrets are slowly being
revealed. Nevertheless, time may be of the essence.
The implications of the undeveloped flowers observed
this year are far-reaching. If plants take several years to
produce flowering stems, individual flowers also take
time to develop, and having several flowers is optimal
for pollination success, then both the rotation age of a
stand and stand structures which produce vigorous,
188
healthy plants are important to the conservation of the

species.
Elliman and Dalton (1995) point out that

preserving large core populations and foregoing the
small, isolated populations is a strategy that is being
adopted by resource managers in some areas. This
management strategy may be appropriate for species
that display a metapopulation dynamic wherein small
populations come and go rather quickly; however,
current information on Cypripedium fasciculatum suggests
the species is adapted to a much longer time scale.
Penelope A. Latham, CFER, Department of Forest Science,

Oregon State University, Corvallis, Oregon 97331
189
TABLES
Table 1. Reproductive characteristics by flower number per stem.
Flower Stems with Fruit

Number Fruits (%) Production (%)
1 per stem 14.7 14.7

2 per stem 54.5 36.4
3 per stem 60.0 36.7
4 per stem 50.0 25.0
Table 2. Reproductive characteristics by site.

Site Total Sample Flowering
Fruiting (%) Stems with >1
Population Population Stems (%) Flower (%)
Site 1 264 57 47.4 49.0
29.8
Site 2 200+ 45 73.3 14.6
22.2
Site 3 110+ 48 14.6 42.8
2.1
Site 4 15 13 84.6 8.3
63.6
190
LITERATURE CITED
Catling, P. M., and V. R. Catling. 1991. A synopsis of breeding
systems and pollination in North American orchids.
Lindleyana 6(4): 187-210.
Cribb, P. 1997. The genus Cypripedium. Timber Press, Inc.,
Portland, OR.
Curtis, J. T. 1943. Germination and seedling development in five
species of Cypripedium L. American Journal of Botany 30: 199-
206.
Elliman, T., and A. Dalton. 1995. Cypripedium fasciculatum Kellogg
ex Watson in Montana. North American Native Orchid
Journal 1: 59-73.
Kunin, W. E., and K. J. Gaston (eds.). 1997. The biology of rarity:
Causes and consequences of rare-common differences. Chapman &
Hall, London.
Rabinowitz, D. 1981. Seven forms of rarity. Pp. 205-217 in The
Biological Aspects of Rare Plant Conservation (ed. H. Synge),
John Wiley and Sons, New York.
Rathcke, B., and E. P. Lacey. 1985. Phenological patterns of
terrestrial plants. Annual Review of Ecological Systems 16:
179-214.
USDA Forest Service, and Department of Interior, Bureau of
Land Management. 1994. Record of decision for amendments to
Forest Service and Bureau of Land Management planning
documents within the range of the Northern Spotted Owl and
standards and guidelines for management of habitat for late-
successional and old-growth forest related species within the range of
the Northern Spotted Owl. Portland, OR.
Whittaker, R. 1960. Vegetation of the Siskiyou Mountains,
Oregon and California. Ecological Monographs 30: 279-338.
191
Plate 1.McCartney
above:
Michaux's orchid
Habenaria quinqueseta
below: night fragrant epidendrum
Epidendrum nocturnum
Collier Co., FL C.L. McCartney
192
Plate 2. Sheviak
Figure2:
Evident
hybrid from
the same
population
as the plants
in Figure 1.
One plant
(collected as
Sheviak
2275b
[NYS])
under
cultivation:
left, first year
after
collection;
right, second
year.
Photos not
to same
scale.
Figure 1: Evident hybrids from a mixed population of P. dilatata var. albiflora and P.
purpurascens in Boulder Co., Colorado. Morphologically these agree with the plant
figured by Luer as P. hyperborea var. gracilis. Note the variation in color between the
putative parents and the tendency for some plants to bear flowers in fascicles.
Sheviak 5896-5897 [NYS]. Photos by C. J. Sheviak
193
Plate 3. Brown
above right:
S. lanceolata var. lanceolata
left: forma folsomii
center: forma lanceolata
right: forma albidaviridis
Folsom's golden ladies'-tresses

Sacoila lanceolata var. lanceolata forma folsomii
Florida's Turnpike P.M. Brown
194
Plate 4. Latham
clustered lady's-slipper
Cypripedium fasciculatum
OR P. Latham
195
LOOKING FORWARD
September 1999
more
Proceedings from the
th
4 Annual North American Native Orchid Conference
including
Orchids of South Forida
Florida - the Rules are Different Here!
An Update on the Symbiotic Seed

Germination of Encyclia tampensis,
Platanthera leucophaea, Platanthera integra
and Spiranthes ovalis var. erostellata
History of the Fakahatchee Strand
The Theory and Practice of Classification
and more!
196

June 1999 North American Native Orchid Journal

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NORTH AMERICAN

NATIVE ORCHID JOURNAL

The Journal welcomes articles, of any length, of both a scientific

2000 Membership in the North American Native Orchid Alliance,

With this the June issue for 1999 I trust the

The September issue will be printed and assembled in

Paul Martin Brown

after September 15:

THE UNPREDICTABLE WEATHER

The Slow Empiricist

It seems that the weather patterns change almost

My first year in retirement in Florida in 1997 was

summer, which produced many wild fires that burned

How has this impacted the world of native

The prolonged dry spell and the warmer than

adder's-mouth, Malaxis unifolia, which was abundant in

Paul Martin Brown, who organized the

Just the week before, in preparation for the

scheduled field trip as a last minute check, they had all

Weather has certainly played a major factor in

My best advice is to persist in your quests and

small spreading pogonia

4TH ANNUAL NORTH AMERICAN

Although the University of South Florida in Tampa,

John Beckner of the Orchid Identification

Roger Hammer of the Dade County Park Service

Larry Zettler, a biologist at Illinois College,

On Sunday, Eric Christenson, a Ph.D. research

show." Wendell Vaught and Mike Owen, both of the

Two members presented some of there favorites

Ken Cameron of the New York Botanical

The concluding speaker was Robert Dressler who

Mike Owen, biologist for the Fakahatchee Strand

Conservation & Education Award for his meticulous

Two days of field trips before the two-day

Back in Florida, near Middleburg and Starke we

National Forest. A hand lens made the difference on

The excessive heat of an early Spring knocked

In the Disney Preserve near Kissimmee, we

At Highlands Hammock State Park, we saw the

superior to the flowers. A big disappointment for me

Our last day, after a brief stop at the airport by

Philip E. Keenan, 31 Hillcrest Dr., Dover, NH. 03820

ORCHIDS AT RABENAU CAMP

In March of 1979, while on a botanical tour in

We got to be friends, and in September of that

The word "camp" made me envision rather crude

But even better than the accommodations was the

communities that included pinelands, open glades,

Although this central camp encompassed only 90

Rabenau Camp is in the northeastern Big Cypress

In the company of Bill's son, William (who was,

In the nearly 20 years that I have been botanizing

encountered 29 species of orchids. All of these species

Of the 29 species of orchids discovered in the

The following is a quick look at the orchids

Epidendrum anceps: A fairly common species of

Epidendrum difforme: Rare in this section

Epidendrum nocturnum: Relatively common in

the entrance to the tube formed by the adnation of the

Epidendrum rigidum: A sprawling rhizomatous

with a purplish-red spot or stripes in the middle of the

Encyclia cochleata: Regularly encountered in